Академический Документы
Профессиональный Документы
Культура Документы
in Rice
1996
NDUAT, IRRI
Suggested Citation :
Singh, V.P. et al. (eds) 1996. Physiology of Stress Tolerance in Rice : Proceedings of the
International Conference on Stress Physiology of Rice, 28 Feb - 5 March 1994, Lucknow,
U.P., India, 239 p.
CONTENTS
Sl.
No.
Particulars
Author (s)
Page
Nos.
FOREWORD
ACKNOWLEDGEMENT
INTRODUCTION
1-10
1.
K.S. Fischer
2.
11-30
3.
31-43
4.
44-69
5.
M.C. Drew
70-79
6.
Response of flooding in
gogorancah rice and moisture
stress effect at reproductive
stage in walik jerami rice.
80-90
S1.
No.
Particulars
Author (s)
Page
Nos.
7.
8.
103-122
9.
123-149
10.
150-167
11.
168-185
12.
186-197
13.
198-206
14.
15.
91-102
207-220
221-239
FOREWORD
The needs of the rainfed lowland rice ecosystem in Southeast Asia are
enormous. The hundreds of millions of the worlds poorest people living in this
region depend upon the almost 40 million hectares of rainfed lowland rice. The
extent of the area combined with a difficult, heterogeneous, and variable
environment demands that a new research approach be adopted. The Rainfed
Lowland Rice Research Consortium was established in 1991 to provide a framework
within which national and international institutes could bring their resources to bear
on this important, though difficult environment. National programs from
Bangladesh, India, Indonesia, the Philippines, and Thailand joined with IRRI, and
supported by the Asian Development Bank, to develop a coordinated research
strategy and program directed towards improving the sustainable productivity of
this environment.
This conference on stress physiology represents several important steps in the
growth of the Consortium. First, it is directed towards assessing the status of strategic
understanding of two of the most critical constraints in the ecosystem: drought and
submergence. Research investment has been increasing in these areas, yet there are
few fora in which scientists can exchange views and findings while focussing on
this ecosystem. A systematic compilation of the present status is essential to careful
planning of future steps.
Second, scientists from Consortium member countries were able to interact
with scientists from advanced research laboratories from Europe, North America
and Australia and begin to plan joint "North-South" research and collaboration.
Scientists from other countries with important rainfed lowland production areas
were able to attend, even though their institutional strengths do not yet permit full
Consortium research responsibilities. It is expected that the contacts made at this
and other similar conferences will stimulate "South-South" collaboration.
Third, lively and vigorous discussions were carried on throughout the event;
both in plenary sessions and in the corridors. The active participation of members
from all the countries and institutions represented indicated that the spirit of
scientific research has permeated the culture of the Consortium. This collegial spirit
is essential if true partnership is to guide research in this complex ecosystem.
Finally, the event was hosted by one of the member institutes, Narendra Dev
University of Agriculture and Technology, with support from the Indian Council of
Agricultural Research, and the Uttar Pradesh Council of Agricultural Research.
This marks an important step in assuming the full range of responsibilities
that come with Consortium membership. It is fitting that this event took place in
the midst of such an important rainfed lowland rice growing area. That the opening
was graced with the attendance of the Governor of the State of Uttar Pradesh is an
indication of the importance that local governments place on improving their
agricultural productivity. Indeed, if Uttar Pradesh were a separate country, it would
be the 7th most populous nation in the world.
I congratulate the participants and the organizers for realizing this event.
These proceedings will serve as a valuable reference for scientists working in this
dynamic field. It is especially significant that, through the Consortium, this product
will be readily available to scientists working in developing countries where access
to the latest physiological literature is severely limited. The host country and the
host institution have made this event a very memorable one. The scientific
competence and a specially warm hospitality has impressed each and every
participant. I am pleased that IRRI working with Consortium members has had the
opportunity to contribute to this important event. I sincerely hope that there will be
many more in this tradition in the future, and that IRRIs role will continue to evolve
towards that of an equal participant among colleagues sharing a common goal.
Klaus J. Lampe
Director General, IRRI
12 September, 1994
vi
ACKNOWLEDGEMENT
vii
INTRODUCTION
The rainfed lowland rice ecosystem covers half the rice growing area in the
humid and subhumid tropics of Southeast Asia. Including the contiguous
flood-prone areas, almost 50 million hectares are sown to rainfed lowland rice. The
system is characterized by environmental heterogeneity and variability, with
frequent droughts and floods damaging the rice crops. Modern rice technology has
barely contributed to rice production and, consequently, yields remain low, in many
areas less than 2 tons per hectare. None the less, this ecosystem provides the staple
food for hundreds of millions of the worlds poorest urban and rural dwellers. But,
as populations and economies continue to grow, increasing pressure is placed upon
the rainfed lowlands to help meet the estimated 70% increase in rice demand
projected over the next 30 years. This pressure is even more acute as investment in
irrigation declines, and water and land are diverted from irrigated rice agriculture
to urban and industrial uses.
Research is required to increase the productivity of this vast ecosystem, and
this increase must be sustainable. That is, the sometimes conflicting demands of
social and economic development must be reconciled with husbandry of our social
and natural resources. Developing technologies that permit such productivity gains
while preserving or enhancing the resource base will demand a fundamental
understanding of the processes that detrermine plant adaptation to the environment,
the dynamics of water and nutrient flow through the system, and the interaction of
management practices designed to maximize efficiencies with the biotic, social, and
economic environment.
The distribution and enormity of the rainfed lowland ecosystem combined
with the complexity of the environment and the difficulties of the production
constraint required that a new research approach be adopted. In 1991 the Rainfed
Lowland Rice Research Consortium was formed. It is a formal linkage among
institutions in five countries (Bangladesh, India, Indonesia, Philippines and
Thailand) and the International Rice Research Institute. The member institutions
agreed upon a research strategy specific to achieving sustainable productivity
increases in the ecosystem, and apportioned responsibilities according to
comparative advantages and available infrastructure and personnel. The Rainfed
Lowland Rice Research Consortium certainly represents a paradigm shift from a
target of uniformly favourable environments addressed from a centralized research
viii
There is little doubt that research into the physiology of stress tolerance can
make a major contribution to improving sustainable productivity of rainfed lowland
environments. However, with the finite resources available to us it is essential to
sharply focus our efforts and take full advantage of the human resources available
to us. Fragmentation of effort and a diffuse approach along more academic lines will
simply delay our progress and frustrate our efforts. This conference has been an
important step in developing the team approach and, if followed up by appropriate
planning and training, should lead to rapid progress over the coming years.
Editors
xi
SUMMARY
Inspite of the vast information on response to water deficits at the crop, and
molecular level, there are few examples of designed change in crop performance
through genetic improvement. This paper suggests that the lack of this knowledge
is partly due to lack of research priority focus and organization of interdisciplinary
teams. The case is made for a sequence of research activities broadly grouped under :
1) measurement of the resource and of the crop response in the components
of resource capture (transpiration), resource use (water-use efficiency) and resource
allocation (harvest index) to define targets for better focus; 2) study of mechanisms
to identify putative traits and develop an "internal consistency" in the relationship
between the traits and field performance; 3) design models at geographic, crop,
genotype (ideotype), and gene level to both priority research opportunities and to
enhance the efficiency of the improvement process; and 4) develop methods to
incorporate selection for adaptation to moisture deficit in routine plant breeding
programs.
This framework, to move from understanding to manipulation, is based on a
case study for the improvement of tropical maize ( Zea mays L.) to moisture deficits.
In tropical maize, intermittent periods of moisture stress are estimated to cause yield
losses of 15% in more than 32 million ha. Maize is particularly vulnerable around
flowering, causing a loss in grain number and through harvest index, grain yield.
Based on this understanding of the mechanism for adaptation of maize to drought,
a "drought plant ideotype" was developed. Eight cycles of recurrent selections for
the ideotype resulted in a gain of 500-800 kg/ha over a range of water-limited
environments. Yield gains were associated with increased ear growth rate around
flowering and harvest index at maturity (resource allocation).
INTRODUCTION
This paper does not review the wealth of information on rice adaptation to
variable and moisture-limiting environments. Instead it provides a conceptual
framework that will allow understanding of the physiology of adaptation to be used
effectively to manipulate the crop and the environment for enhanced productivity.
1 The International Rice Resarch Institute, P.O. Box 933, Manila, Philippines.
From the plant perspective, the crop in the field is a community of genetically,
fairly uniform plants representing the "genotype". During a growing season the
particular combination of soil, climate, and pests, plus the competition among crop
plants and other species (weeds) forms a unique environment, which when
interacting with the "genotype", leads to the expression of the phenotype, and
ultimately yield.
Much of our success in improving rice productivity was the result of enhancing
and stabilizing the environment, particularly for water (irrigation) and nutrients
(fertilizer) and at the same time increasing the genotype's ability to respond to this
change in the environment with increased yield. There was an additional benefit.
Irrigation and nutrients reduced the number of unique environments determined
mainly by temperature ranges" and the occurrence of pests. Fewer unique
environments required fewer unique genotypes to maximize the phenotypic
expression. We were able to develop rice cultivars adapted over a broad geographic
range and responsive to a given and predictable environment.
In contrast, the rainfed lowlands have a range of environments whose
diversity is driven by variability in, for example, the amount, intensity, and duration
of water. Each of the key environmental variables, eg., moisture regime, fertility,
flooding incidence, biota, daylength is complex in itself and interacts with the others
and with the rice plant. The resultant phenotype-the expression of the genotype in
these variable environments-is by way of the metabolic processes of the plant, i.e.,
its physiology. Given the environmental complexity, there are practically an infinite
number of expressions of phenotype. Thus, to advance crop production in these
environments through directed improvements in phenotype, those growth processes
that are interacting, rather than the myriad of interaction themselves must be
understood. That is, understanding leading to the rice phenotype is one means to
reduce enormous complexity to a manageable level.
Wilson (1992) contends that formal treatment of physiology gave information
as to how the crop grows, but little on how it might be manipulated genetically to
achieve particular results. This paper examines some case studies in other cereals
(but mainly maize) where such manipulations have occurred. Passioura (1981)
contends that breeding the changing of a genotype-is a practical affair with its
successes and failures being judged by the farmer clients. In contrast, the success of
physiology understanding the growth processes of the plant-depends on how
enlightening it is in providing ideas and tools for the breeding practitioners.
Our task is to develop realistic means to evaluate the contribution of
physiology research. Thus a set of "signposts" that will generate realistic ideas and
understanding, which then become appropriate tools for manipulation by the
practical breeder is proposed. These are measurements, mechanisms, models, and
methods.
SIGNPOSTS
Measurement
What to measure and why? When the new science of crop physiology began
some 40 yr ago, there was an expectation that simply understanding and measuring
such processes as photosynthesis and transpiration, would provide a direct benefit
to yield. In retrospect, we began to look at very fine-albeit critical- processes at the
very beginning without generating a broad conceptual framework of crop
development. The first measurement needed considers the basic components of the
limiting resource and how it is used, viz. resource capture, conversion and
partitioning. Under water-limiting environments, grain yield is a function of the
water transpired (T), the water-use efficiency (WUE) and the partitioning of dry
matter to grain (HI). One or more of these components must be altered to increase
grain yield:
Thus, before embarking on a breeding program for improvement under
variable and water-scarce environments, one must first measure how the current
system uses the scarce resource (water) and where the opportunities for genetic
intervention lie. We need to measure how much of the total precipitation is actually
transpired and how much is lost in runoff, soil evaporation and drainage, and how
much is stored in the soil. We must also know if there is variation in WUE and HI
(Fig. 1).
Fig. 1.
A scheme of water balance showing stored soil water (S) as a target for improvement of
sorghum under terminal drought.
Let us consider two case studies-maize and sorghum (Sorghum bicolor (L.).
In maize, Kassam et al. (1975), the pioneers of todays modelers fitted the water
requirements of maize to the bimodal rainfall patterns of the savannas in West Africa.
Their model predicted two opportunities for improving the performance of maize
in this environment (Fig. 2). In the first season, there was a high probability that
demand for water for transpiration will exceed supply midway during
growth-around flowering. Studies (Shaw, 1977; Westgate and Bassetti, 1990; Boyer,
1992) have shown flowering to be the most sensitive stage of growth in maize;
moisture stress at flowering severely reduces grain number and harvest index (HI).
Thus one target for the improvement of maize grown under variable and
unpredictable moisture supply is to modify flowering behavior and grain number
and thereby increase yield through higher HI. Ed meades et al. (1993) have estimated
that more than 32 million ha of maize grown in the lowland tropics are exposed to
intermittent drought around flowering causing 15% losses in yield.
Fig. 2
In these two examples, measurement of the limiting resources and its effect
on yield component set clear targets and focus for crop improvement. Setting such
targets is critical in the rainfed ecosystem. Because of the variability (and large
genotype environment interaction), the selected targets will not be expressed in
all locations at all times. Careful measurement allows for identifying the repeatable
and useful positive genotype environment interaction which will have greatest
benefit over time at any one location.
Mechanisms
Thew is a need to understand the mechanisms and the processes that will
increase either resoure capture, resource-use efficiency or allocation.
In the past, physiologists were content to show a correlation between a given
plant character and yield-without differentiating causality from spurious, although
statistically significant correlation. A good example of this misuse is the attempt to
improve grain yield by selecting for either an increase in grain number or grain size.
This is bound to be ineffective if grain yield is limited by the capacity to produce
assimilates for grain storage and not by the storage system of the grain itself. Not
only must the presence, strength and direction of a process be correlated with the
targeted yield component there also must be physiological "internal consistency" in
each of the intervening steps.
In sorghum, a target is to increase root exploration for soil moisture at depths
during grain filling. One mechanism for that is osmotic adjustment (OA)- the change
in osmotic potential due to the accumulation of solutes within the cell. Osmotic
adjustment maintains turgor pressure under moisture stress and thus root activity.
It also increases dehydration tolerance and in terminal stress, enhances the
retranslocation of assimilates to the grain (Santa maria et al., 1986). It is important to
demonstrate that OA is not just related to grain yield, but also that there is an internal
physiological consistency whereby
*
Increased root growth delays leaf area loss under moisture stress and
increases dry matter; and
Fig. 3
Effect on maize grain yield from 1d of moisture stress (a) and 1d with 54% crop shading (b).
A characteristic of maize under environmental stress is increased anthesis-silkinginterval. The reserves of carbohydrate in the maize plant at silking are comparatively
small and the capacity of newly fertilized ovules to attract these reserves is limited.
Boyle et al. (1991) infused the stems with a sucrose-rich solution to offset most of
the effects of drought stress at flowering. Edmeades et al. (1993) concluded that the
reduced grain number per plant under drought seems more of a chemical (CHO
supply) than physical (separation of pollen and stigma) process, and that the
anthesis-silking interval is a symptom of reduced assimilate flux rather than the
direct cause of barrenness. Here the internal physiological consistency links the
relationship between anthesis-silking interval and yield (Fig.4), with reduced
assimilate supplies around flowering for ovule and silk growth (Fig.5) and to reduce
grain number and HI.
Fig. 4
Relationship between anthesis silking interval and grain yield combined data from maize
cultivars grown under drought (o) and well-watered conditions () at Tlaltizapan, Mexico.
From Bolanos and Edmeades (1993).
Fig. 5
Relationship between mean ear spikelet biomass at 50% anthesis and the anthesisto-silking (ASI) interval for Tuxpeno Sequia cycles 0,2,4,6 and 8 grown under high plant
density and mild severe drought stresses.
Models
Modelling processes and determining the likely effects of changes in plant
characteristics aid in the development of "best bet" approaches. Processes that cannot
be modelled are probably not understood. Without this understanding, our move
from measurement to new technologies will be unpredictable (and quite often
unsuccessful).
There are many levels of modelling-they do not have to be complex if that is
not necessary for the job at hand, The conceptual model or the "ideotype" has served
physiologists and breeders well. The most successful dates from 1960, with the IR8
story of IRRI. In 1974 Jennings, the IRRI breeder, involved in the green revolution
said:
"elegant work by the institute crop physiologists indicated in
detail on paper an ideal plant type, i.e., an "ideotype", and pinpointed
the vegetative characteristics of traditional varieties requiring drastic
modification. In retrospect it was a relatively simple task for breeders
to follow this blue print and produce a series of productive dwarfs
beginning with IR8".
Fischer et al. (1983) and later refined by Edmeades et al. (1993) developed an
ideotype for the selection of maize for better performance under limiting moisture
around flowering. They used this information in a selection index for a field site
recurrent selection program for the improvement of maize performance under
moisture deficits.
7
The ideotype concept which creates scientific ideas, and a framework within
which they can be tested should remain a strong tool for a breeding program. The
latest application is in the development of a new plant type to raise the yield frontier
in tropical rice. Originally, almost as an intuitive tool, our latest ideotypes have
involved crop modelers and physiologists from the outset. We have advanced in our
understanding of processes since the 1960s and now it is possible not only to develop
the conceptual model (as for IR8), but also process-driven models with which
sensitivity analysis-the "what if" questions- can be done in the computer before going
into the field (Kropff et al., 1994).
Models at the crop and molecular levels are now enhancing our efficiency in
research. Recent applications of process-based rice crop models linked with GIS and
to reliable data bases are allowing targeting ideotypes and assessing the performance
of alternative crop ideotypes. At the molecular level, we are developing the genetic
maps for the putative traits of root length, root penetration, osmotic adjustment and
desiccation tolerance as "models"to aid the selection process. This work is reported
in more detail elsewhere in these proceedings. And just as for past studies at the
process level, we must ensure that our molecular maps are developed from
appropriate populations to ensure that the genetic information is sufficiently robust
for application for selection within the agronomically fit population growing in the
breeder's field.
Method
This is the most critical of the signposts, requiring physiologists and breeders
to work together to be productive. The synthesis of the previous steps must come
here, or the effort invested will serve merely academic interests.
In the past, physiologists have tended to disregard genetic variation.
Extrapolation and generalization were often derived from a few, or worse, one
genotype. Correlations which depend on one extreme outrider entry were selected
from unadapted material from a germplasm bank. Too often a relationship among
yield-determining characters was established in controlled environments and never
in the field. Indeed, physiologists have worked downstream from the breeding
program spending time telling the breeders what they managed to accomplish, rather
than assisting in the current selection process.
This has led to charges that physiology is a retrospective science. The guiding
rules to applying our knowledge and skills should include (a) that physiologists
work with many genotypes representative of the variation in breders fields; and
(b) that the physiologists maintain a well-worn path to those breeders' fields.
The maize, ideotype was developed from the study of mechanisms in a broad
range of maize germplasm and from an analysis of the variation that exist within
one maize population (Tuxpeno), which is used as a breeding population for new
varieties (Fischer et al., 1983).
Using the ideotype characters for selection, divergent subpopulations were
developed and their performance compared with that of subpopulations selected
8
on yield alone. The study demonstrated that a) genetic variation for drought
characteristics existed within agronomically fit populations being used in breeding
programs; b) the ideotype selection enhanced performance under yield-limiting
moisture around flowering; c) there was an internal consistency linking change in
the population through selection to the processes that influence adaptation to
moisture deficits; d) selection for the drought ideotype was more effective than
selection for yield alone; and e) screening for the additional traits was practical (although
costly) for a breeding program.
IMPACT
Recently Edmeades et al. (1993) have reported progress after six selection
cycles for maize improvement to the target drought environment. The evaluation
studies included entries from the same breeding population where selection was for
yield (alone) done over a number of test locations (the more conventional approach
for breeding). Selection for drought tolerance resulted in a significant increase in
grain yield at levels ranging from 1 to 8 t/ha, with no significant interaction between
rate of gain and yield level. At 2 t/ha (common under drought conditions) this
genetic improvement represents a gain of 6.3% per cycle.
Gains in yield were the result of reduced barrenness under moisture dificit at
flowering. Total biomass under drought was unaffected by selection, so the gains in
grain yield arose from a systematic increase in harvest index-the target component
of selection. This was accompanied by an increase in ear biomass at 50% anthesis
and a decrease in the number of tassel branches (a change in the allocation of
resources). Flowering behavior was markedly affected by selection, with a large
decrease in the ASI, which was related to ear spikelet biomass at flowering
(Edmeades et al., 1993).
Based on this study, selection for adaptation to moisture stress around
flowering is being practiced in a number of maize populations as a routine in the
breeding program of CIMMYT.
REFERENCES
Bolanos J, Edmeades G O (1993) Eight cycles of selection for drought tolerance in lowland tropical maize.
I. Responses in grain yield, biomass, and radiation utilization. Field Crops Res. 31:233-252.
Boyer J S (1992) Mechanism for obtaining water use efficiency and drought resistance. Pages 181-200 In:
Plant breeding in the 1990's. H.T. Stalker and J.P. Murphy, eds. Commonwealth Agricultural Bureaux
International, Wallingford, United Kingdom.
Boyle M G, Boyer J S, Morgan P W (1991) Stem infusion of liquid culture medium prevents reproductive
failure of maize at low water potential. Crop Sci. 31:1246-1252.
Edmeades G O, Bolanos J, Hernandez M, Bello S (1993) Causes for silk delay in a lowland tropical maize
population. Crop Sci. 33:1029-1035.
Fischer K S, Johnson E C, Edmeades G O (1983) Breeding and selection for drought resistance in tropical
maize. El Batan Mexico.
Grant R F, Jackson B S, Kiniry J R, Arkin G F (1989) Water deficit timing effects on yield components in
maize. Agron. 1. 81:61-65.
Kassam A H, Kowal J, Dagg M, Harrison M N (1975) Maize in West Africa and its potential in Savanna
areas. World Crops 27:73-78.
Kropff M J, Cassman K G, van Laar H H (1994) Improving yield potential: the role of ecophysiological
crop models. In : Proceedings of the International Rice Research Conference, 1992 IRRI, Los Banos,
Philippines.
Passioura J R (1981) The interaction between the physiology and the breeding of wheat. Pages 191-200.
In : Wheat science-today and tomorrow. LT Evans & WJ Peacock, eds. Univ. Press, Cambridge, London.
Santamaria F J, Ludlow M M, Fukai S (1986) Drought resistance traits in hybrids and lines of Sorghum
bicolor L Moench. Pages 127-143. In: Proceedings of the first Australian Sorghum Conference, M.A. Foale
and R.G. Henzell, eds .
Shaw R H (1977) Climatic requirement: Corn and Corn Improvement. G.F. Sprague, ed. Am. Soc Agron.
18:591-623.
Westgate M E, Bassetti P (1990) Heat and drought stress in corn: what really happens to the corn at
pollination? Pages 12-28. In: Proceedings of the 45th Annual Corn and Sorghum Research Conference,
5-6 Dec 1990. D. Wilkinson, ed. American Seed Trade Association Washington, DC, USA.
Williams R L (1990) Genotypic variation in delayed leaf senescence in sorghum. MS Thesis Univerisity
of Queensland Australia.
Williams T V, Snell RS, Ellis JF (1976). Methods of measuring drought tolerance in corn. Crop Sci. 7:179-181.
Wilson G L(1992). Crop physiology: Some recollections and current perceptions. Pages 2-10 In: Proceedings
of the 6th Australian Society of Agronomy Conference, Armidale, Australia.
10
SUMMARY
Submergence of rice is a complex phenomenon acting mainly via the 104 -fold
slower diffusion of gases in solution than in the gas phase. This will lead to both
increases and decreases of supply of oxygen and CO2 and increases in the
concentration of the phytohormone ethylene. This paper focuses on possible O2
deficiencies; without claiming that O2 deficiency is more important than other
possible adverse effects of submergence. The occurrence of O 2 deficiency in
submerged rice is reasonably certain. However, we do not know whether such O2
deficiency will affect survival, or at least long-term growth and yield of the crop.
Low O2 concentration occurs in the water of flooded rice fields, particularly
at greater water depths and also during the night. Available evidence suggests that
O2 deficiency occurs, in partially or wholly submerged rice plants, at least at certain
hours of the day and in certain regions. During the night, O2 concentration at the
epidermis of the root tips is zero and the roots synthesize ethanol, the end product
of anaerobic catabolism. All this ethanol is consumed during the first hours of the
light period. This suggests the possibility that ethanol may be used as a source of
carbon, both in submerged rice and in nonsubmerged plants of paddy fields.
Armstrong's model indicates that O2 deficiency in roots of vascular plants is
particularly likely in the stele of the root tips; O2 supply from the environment to
this core being limited by diffusion and O2 uptake along the pathway. We review
metabolic evidence for roots which is consistent with these models, no information
being available on submerged leaves.
In reviewing mechanisms of anoxia tolerance, we assume universal
requirements for retention of membrane integrity despite a greatly reduced
energy supply and reductions in requirements of energy for maintenance. Using
these requirements as a prerequisite, we present evidence that there are two
different modes of adaptation to anoxia, based on (i) rapid and (ii) slow rates of
catabolism, respectively. We also suggest anoxia tolerant plants with the mode
of slow rates of catabolism may be particularly useful during transient O 2
deficiency as often occurs in floodprone rainfed environments.
Injury due to exposure to anoxia can also be aggravated upon return from
anoxia to aerated conditions, for example production of free radicals of oxygen
would aggravate membrane damage and injury in several species. Whether this
occurs in rice is unknown.
The response to anoxia, which involves a large-reduction in energy production
is also relevant to other situations, for example to carbohydrate starvation, which
can in principle occur due to other factors of a submerged environment such as low
CO2 supply and low light.
INTRODUCTION
The complex environment in flooded rice fields is shown in Fig. 1. The 104 -fold
slower diffusion of gases in solution than in the gas phase (Armstrong, 1979) means
the solution immediately outside the shoot epidermis can contain high O2
concentrations during the day and low concentrations during the night. Theopposite
holds for CO2 concentrations. This conclusion receives support for O 2 from data by
Setter et al. (1988a) for gas concentrations in floodwater in rice fields in Thailand.
These studies also showed that floodwater contained high concentrations of the
phytohormone ethylene (Setter et al., 1988b).
The rest of this review will be confined to the posssible occurrence of O2
deficiency in submerged rice and if so what mechanisms of adaptation may be
required. However, it should be pointed out that one requirement for tolerance to
anoxia, ability to survive at a low level of energy production, is also relevant to cases
Fig. 1.
12
Diagram showing the large range of adverse factors which may occur during submergence
of rice (Setter et al., 1993).
of carbohydrate starvation, whether these are due to low CO2 low light, or chlorosis
caused by high ethylene concentrations, which develops in a submergence intolerant
cultivar (Jackson et al., 1987).
Occurrence of O 2 deficiency depends on :
1)
2)
i)
ii)
iii)
Low metabolic activity coupled with survival, i.e. by the tissue going
into suspended animation, thus conserving carbohydrates for recovery
after the floodwaters recede.
2)
Possible "post anoxic injury" due to return to aerated conditions has also to
be considered. Transfer of tissues from anoxia to air, i.e. restoring the oxygen supply.
Introduction of O 2 may lead to formation of free radicals of oxygen and further
membrane damage, particularly when there has been deterioration of membrane
components during the O2 deficiency itself.
Finally, the relevance of the present paper is for the rice ecosystem rather than
for rice per se : the rice weed, barnyard grass, is probably even more efficient than
rice growing under anoxic conditions. Understanding the mechanisms of the
13
adaptation of barnyard grass could lead to more efficient methods of its control
(Kennedy, et al., 1993).
Definition of Terms
anaerobic catabolism : breakdown of sugars to supply energy in the absence of air.
anaerobic proteins: proteins induced by O2 deficiency.
anoxia: absence of oxygen.
anoxic core: centre of a tissue which does not receive oxygen.
ATP: adenosine trios-phosphate.
daltons: a measure of molecular size similar in magnitude to molecular weight.
glycolysis: breakdown of sugars to the three carbon compound pyruvate.
hypoxia: low but not zero oxygen.
Km: substrate concentration at which half the maximum velocity of a reaction is
attained.
maintenance requirement: energy required without any increase in cell substances.
NAD: nicotinamide adenine dinucleotide.
O2 deficiency: Plant tissues can not obtain sufficient oxygen for maximum respiration
post anoxia: after return from anoxia to air.
PPi : pyrophosphate.
RNA: ribonucleic acid.
DETAILED REVIEW
Occurrence of O2 deficiency in the field
Low O2 concentrations have been shown at greater depth and during the
night in floodwaters of fields of floating rice in Thailand (Setter et al., 1987, 1988a)
and India (Setter et al., this proceedings). O2 concentrations in the floodwaters
decreased with water depth, while in the surface layers of water O2 concentrations
increased during the day and decreased during the night. O2 concentrations inside
floating rice plants are only available for the lacunae, and these show the expected
decrease in O2 concentrations with increasing distance from the O2 supply, i.e. with
increasing water depth (Setter et al., 1987). Similar patterns of O2 distribution in roots
of paddy rice were established in the pioneering studies by van Raalte (1940) in the
Botanical gardens of Bogor. Such patterns are predicted by the elegant models on
O2 distribution within mots of vascular plants, depending on O2 supply via the
aerenchyma (Armstrong and Beckctt, 1987).
No such models for O2 supply exist yet for submerged plants, nevertheless
we can use the model of Armstrong and Beckett to predict that : (i) during the night,
14
O2 supply to roots of submerged rice plants will be substantially lower for submerged
than for non-submerged plants: with O2 close to the shoot root junction at 7% for
the submerged plants (Setter et al., 1987) and an expected 20% O2 in the gas spaces
of non-submerged plants(Armstrong and Beckett, 1983, (ii) O2 supply is likely to
be less for tissue away from the principal O2 supply, i.e. the lacunae or aerenchyma.
As one example, in deep water rice O2 deficiency may become acute in aquatic roots
in deep water unless their epidermis has a low permeability to O2 .
Different modes of response by vascular plants to O2 deficiency
We distinguish four different modes of response of plants to low O 2
concentrations in at least part of their environment.
Mode 1 : intolerant to waterlogging or submergence. These plants have both low porosity
and low cellular tolerance to anoxia and comprise the bulk of plant species.
Mode 2: tolerant to waterlogging and submergence based on high porosity of tissues
to facilitate O2 flow from the air or from water containing O2 (Armstrong, 1979, Armstrong
and Beckett, 1987). Some plants with this mode are not necessarily tolerant to anoxia (cf.ap
Rees et al., 1987) and this has been claimed for rice roots (Webb and Armstrong, 1983). We
do not agree and include rice into Mode 4, as discussed below.
Mode 3: tolerance based on cellular tolerance to anoxia. This mode can be further
divided into two types: 3A) where a tissue can grow under anoxia, for example, rice coleoptiles
(Jackson and Drew, 1984; see also Fig.2) and some species of Echinochloa (Kennedy et al.,
1992), 3B) tissue survives, but does not grow until aeration is restored. The best examples
are germinating seeds of several species (to be discussed in section 3).
Fig. 2.
Genotypic differences in anoxia tolerance of rice during germination (Setter et al., 1994).
Conditions : seeds imbibed 1 day air, followed by 3d anoxia or air treatment.
15
Mode 4: a tolerant mode based on a combination of Modes 2 and 3. This mode can
cope with temporal decreases in O2 regime for example during diuranal cycles of O2 supply.
This combined mode also allows for "symbiosis" between anoxic and aerobic regions in the
same tissue. Well known examples are aerenchymous root tissues with anoxic steles and
aerobic cortices (Armstrong and Beckett, 1987). Submerged established rice plants
presumably belong in this group (to be discussed in more detail in Section 5).
Anoxia tolerance and the production and flow of energy under anoxia
We postulate that universal requirements for anoxia tolerance are the
reduction of maintenance requirements for energy and the maintenance of
membrane integrity, Evidence for the second proposition was obtained for beet root
in which the endogenous dye betacyanin is an excellent marker for cell death (Zhang
et al., 1992). Rapid leakage of K+ , C1- and Na + started up to 9h before leakage of
betacyanin indicating increases in membrane permeability caused death, rather than
vice versa (Zhang et al., 1992, Zhang and Greenway, 1993).
Production and flow of energy to processes essential to survival. Generation of
energy during anoxia will occur mainly via glycolysis {equation 1) with ethanol as
the principal end product of anaerobic catabolism in nearly all plants (equation 2 :
Davies, 1980; ap Rees et al., 1987). The overall reactions are:
16
2)
Table 1.
Plant
Carbonanoxia /
Carbon air
Reference
1.8
Calculated by us from
Alpi and Beevers (1983)
3.0-3.7
1.9
<1.0
0.7
The energy produced during glycolysis in anoxic tissues is used for different
processes:
1)
2)
17
3)
Growth. Elongating rice coleoptiles require net cell wall and protein
synthesis (Alpi and Beevers, 1983), as well as uptake of solutes to
generate osmotic pressure (Atwell et al., 1982). The high energy
requirements, particularly for protein synthesis, may account for the
high rate of catabolism found in rice coleoptiles (Table 1).
18
1)
2)
3)
4)
Enzyme
Alcohol dehydrogenase
7 fold
Lactate dehydrogenase
4 fold
Sucrose synthase
3 fold
PPI
7 fold
dependent-phosphofructokinase
seedling may become anoxic. Once O2 supply to a tissue becomes restricted rather
than cut off altogether anoxic and aerobic tissues will coexist in the same organ.
The hypothesis that this combination occurs is based on the very high affinity of
cytochrome oxidase for O2 indicated by a Km of 24 m molm-3 (Yocum and Hackett,
1957). So electron flow and oxidative phosphorylation will continue till very low
O2 concentrations are reached. As shown schematically in Fig. 3, O2 will diffuse
into the tissue, while O 2 consumption will decrease the O2 concentration along
the pathway. In environments of low external O2 concentration the concentration
gradient will be insufficient to deliver O 2 to the core of the root i.e. this core will
become anoxic. Analogous arguments hold for other tissues, such as submerged
leaves and when the O2 supply is the aerenchyma rather than the external
environment.
Fig. 3.
Diagram showing the O 2 profile expected in a root when the O 2 supply is from the nutrient
solution surrounding the roots (adapted from Armstrong and Beckett, 1987).
Anoxic cores in aerenchymous rice mots have been demonstrated using an O 2 micro
electrode (Fig. 4). The radial pattern of O2 concentrations is consistent with earlier
formulated sophisticated models (Armstrong and Beckett, 1987) based on : 1)
porosity of the tissues; 2) O 2 uptake by respiration; 3) radial O 2 loss to the
environment and 4) length of longitudinal pathway. No such models exist for
submerged leaf tissue but these could be developed in cooperation between
biologists and mathematicians.
The data on O2 concentrations are complimented with metabolic evidence
for an anoxic stele and aerobic cortex in maize roots (Thomson and Greenway,
1991), exposed to low but not zero O2 concentrations or depending on O2 supply
20
Higher concentrations of ethanol in the stele than in the cortex (Table 3).
2)
3)
Table 3.
Ethanol (mol m-3 ) in cortex and stele of excised maize mots exposed to
different O2 concentrations at 25C. (Thomson and Greenway, 1991).
stele
external solution
0.06*
0.39
0.12
0.11
0.00
0.34
0.43
0.42
In Table 4, the inactive and active state of PDC in the cortex and stele,
respectively, is shown by: (i) a lag after addition of pyruvate i.e. the enzyme is
gradually activated in the presence of its substrate. In hypoxic roots such lags are
pronounced in the cortex but slight in the stele, ii) differences in activation of the
isolated enzyme when incubated at pH 6.0; these were large for the cortex but small
for the stele of hypoxic roots (Table 4) suggesting most of the PDC is already
activated.
Evidence for occurrence of anoxic regions in submerged, established rice plants :
The following evidence is available :
1)
Fig. 4.
22
O 2 concentrations (A) and growth of root tips (B) during submergence of rice. O2 was
measured with a micro O 2 electrode at the epidermis of the 2-7 mm zone behind the root
apex of submerged rice plants (Waters et al., 1989). Solid bars on the x-axis represent dark
periods.
Table 4.
Cortex
% increase in activity
Stele
Cortex
1.1
1.9
63
68
0.2
1.4
25
50
0.3
0.7
20
23
O2 Treatment
Stele
Lag (min)
* Air saturated
2)
Roots of intact submerged rice plants produced ethanol during the night
and consumed ethanol at the start of the light period (Fig. 5). Mirror image
diurnal cycles were found for O2 with loss from the roots during the day
and O2 uptake during the night (Fig. 5)
Fig. 5.
Ethanol synthesis during night and ethanol consumption during the day by roots of intact
submerged rice plants (Waters et al., 1989). The roots were in a sealed container so that
O2 and ethanol in the nutrient solution surrounding the roots could be measured. Solid bars
on the x-axis represent dark periods.
where they can contribute to the carbon balance. Similarly, compounds able to
contribute to energy supply such as ATP and PPi may be transported from aerobic
to anoxic regions. Presumably such an input would allow much more synthetic
activity than when the entire tissue was anoxic. Most of this argument is mere
speculation and the only evidence available is for ethanol shown in Fig 5 and by
transport of ethanol from waterlogged roots of tomato plants to their shoots (Fulton
and Erickson, 1964).
In this paper we further consider possible ethanol metabolism in the aerobic
regions of an hypoxic tissue without claiming this is the only or most important
exchange of metabolites occurring in tissues with aembic and anoxic regions. Rates
of ethanol consumption are of the same order of magnitude as those of ethanol
formation (Table 5). The observed rate of ethanol consumption by rice mots is 4-fold
lower than the maximum observed rate in peas (Table 5), however, this could be
due to the difference in exogenous ethanol concentration rather than due to species
difference.
During ethanol consumption, acetate is formed together with 2 mols of
NADH/mol of ethanol as observed for pea and mungbean tissues (Cossins, 1978).
The carbon is subsequently incorporated in a large range of metabolites, similar to
carbon derived fmm glycolysis (see Cossins, 1978 for review). The pathway from
ethanol to acetate can continue as long as the tissues are aerated because NAD can
then be regenerated from NADH in several reactions and by conversion to ATP in
oxidative phosphorylation.
24
Table 5.
0.033
0.12
0.042
ethanol
2)
Rates of ethanol
concentration.
3)
4)
5)
6)
consumption
as
related
to
external
ethanol
25
7)
Species
Lipid degradation
products (1)
Free radical
formation (2)
Superoxide
dismutase(3)
Iris pseudacorus
(tolerant)
Few
None observed
Iris germanica
Large number
Rapid genera-
Either decreased
tion
or small increase
Glyceria maxima
(intolerant)
Fig. 6.
CONCLUSION
Anoxia tolerance
We postulate two modes of anoxia tolerance. Both modes require maintenance
of membrane integrity and sophisticated regulation to direct the limited amounts of
energy to processes essential to survival, however, they differ in the rate of
catabolism :
27
i)
ii)
iii)
iv)
Long term anoxia of all plant tissues probably only occurs under
flooded conditions during the early period after direct seeding.
Anoxic regions coexisting with other regions receiving sufficient O2 for
oxidative phosphorylation may be the norm for submerged rice and
may even occur in the roots of paddy rice. At least in submerged rice
the anoxic regions will be more voluminous during the night than
during the day.
The best mode of anoxia tolerance for the association of aerobic and
anoxic tissue is unknown.
It is also unknown whether O 2 deficiency adversely affects submerged
rice plants.
REFERENCES
Alpi A, Beevers H (1983) Effects of O2 concentration on rice seedlings. Plant Physiology 71:30-34.
ap Rees T, Jenkin L E T, Smith AM, Wilson PM (1987) The metabolism of flood tolerant plants. Pages
227-238. In : Plant life in aquatic and amphibious habitats. R.M.M. Crawford.ed. Blackwell Scientific
Publication, Oxford.
Armstrong W (1979) Aeration in higher plants. Pages 226-332 In: Advances in Botanical Research. H.W.W.
Woolhouse, ed. Academic press, London.
Armstrong W, Beckett PM (1987) Internal aeration and the development of stelar anoxia in submerged
roots. New phytologist 105:221-245.
Armstrong W, Cringle S, Brown M, Greenway H (1993) In: Interacting stresses on plants in a changing
climate, NATO advanced workshop AS1 Series 1 Vol 16. M.B. Jackson and C.R. Black, eds. Wye College.
Atwell BJ, Waters I, Greenway H (1982) The effect of oxygen and turbulence on elongation of coleoptiles
of submergence tolerant and submergence intolerant rice cultivars. J.Expt. Rot 33: 1030-1044.
Bailey-Serres J, Kloeckener Gruissem B, Freeling M (1988) Genetic and molecular approaches to the study
of the anaerobic response and tissue specific gene expression in maize. Plant Cell and Environment 11:
352-357.
Cossins E A (1978) Ethanol metabolism. Pages 169-202 In: Plant life in anaerobic environments. D.D. Hook
and R.M.M. Crawford, eds. Ann Arbor Science Publisher Inc, Michigan.
28
Cossins E A, Beevers H (1963) Ethanol metabolism in plant tissues. Plant Physiology 38: 375-380.
Crawfod R M M (1993) Plant survival without Oxygen. Biologist 40: 110-114.
Crawford R M M, Wollenweber-Ratzer B (1992) Influence of L- Ascorbic Acid on Post-anoxic growth and
survival of chickpea seedlings (Cicer arietinum L.) J.Exy. Bot. 43: 703-708.
Darnell J, Lodish H, Baltimore D (1990) Molecular Cell Biology 2nd Edition Scientific American Books
W.H. Freeman and Co. New York.
Davies D D (1980) Anaerobic metabolism and the production of organic acids. Pages 581-611. In: The
Biochemistry of Plants. Vol.2 D.D. Davies. ed. Academic Press, London.
Drew M C (1990) Sensing soil oxygen. Plant Cell and Environment 13: 681-693.
Faiz-ur-Rahman A T M, Trewavas A J, Davies DD (1974) The Pasteur effect in carrot root tissues. Planta
118 : 195-210.
Fulton J M, Erickson A E (1964) Relation between soil aeration and ethyl alcohol accumulation in xylem
exudate of tomatoes. Proc. Soil Sci. Soc of America 28 : 610-614.
Jackson M B, Drew MC (1984) Effects of flooding on growth and metabolism of herbaceous plants. Pages
47-128 In:Flooding and plant growth. T.T. Kozlowski, ed. Academic Press, London.
Jackson M B, Waters I, Setter TL, Greenway H (1987) Injury to rice plants caused by complete submergence:
a contribution by ethylene(Ethene). J.Exp. Bot. 38:1826-1838.
John C D, Greenway H (1976) Alcoholic fermentation and activity of Some enzymes in rice roots under
anaerobiosis. Aust J. Plant Physiol. 3:325-336.
Johnson J, Cobb BG, Drew M C (1989) Hypoxic induction of anoxia tolerance in root tips of Zea maize. Plant
physiology 91:837-841.
Kennedy R A, Rumpho M E, Fox TC (1992) Anaerobic metabolism in plank. Plant Physiology. 100:1-6.
Mayne R G, Lea P J (1985) Properties of three sets of isoenzyme of alcoholdehydrogenase isolated from
barley (Hordeum vulgare). Phytochemistry 24: 1433-1438.
Menegus F, Cattaruzza L, Mattana M, Beffagna N, Ragg E(1991) Response to anoxia in rice and wheat
seedlings. Plant Physiology 95: 760-767.
Mocquot B, Prat C, Mouches C, Pradet A (1981) Effect of anoxia on energy charge and protein synthesis
in rice embryo. Plant Physiology 68:636-640.
Mohanty B, Wilson P M, ap Rees T (1993) Effects of anoxia on growth and carbohydrate metabolism in
suspension cultures of soybean and rice. Phytochemistry 34: 75-82.
Monk L S, Fagerstedt K V, Crawford R M M (1987). Superoxide dismutase as an anaerobic
polypeptide. Plant Physiology 85: 1016-1020.
Monk L S, Fagerstedt K V, Crawford R M M (1989) Oxygen toxicity and superoxide dismutase as an
antioxidant in physiological stress. Physiologia Plantarun 76:456-459.
Penning de Vries F W T (1975) The cost of maintenance processes in plant cells. Annals Rot. 39: 77-92.
Pradet A, Bomsel J L (1978) Energy metabolism in plants under hypoxia and anoxia. Pages 89-118. In: Plant
life in anaerobic environments. D.D.Hook and R.M.M. Crawford, eds. Ann Arbor Science Publisher Inc,
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Raymond P, Pradet A (1980) Stabilization of adenine nucleotide ratios at various values by an oxygen limitation
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Robards A W, Lucas W J (1990) Plasmodesmata. Annu. Rev. of Plant Physiol. Plant Mol. Biol. 41: 369-419.
29
30
INTRODUCTION
Higher plants must be well aerated in order to survive, compete and complete
their life cycle successfully. All plant parts require oxygen for respiration and other
biochemical oxidations/ oxygenations. Besides, green shoots require an external
input of carbon dioxide for photosynthesis that benefits the whole plant. Plants also
need to dispose the potentially active or damaging volatile metabolites such as
photosynthetic oxygen, acetaldehyde, ethylene or methyl jasmonate produced by
various metabolic pathways. The principal environmental factor affecting aeration
is water. Little available water adversly affects the growth whereas, too much water
can asphyxiate and may kill growing seedlings in majority of cases. Slow diffusion
of gas in water compared to air (D water / Dair) is approximately 1.13 10-4 which is
responsible for this traumatic effect of an otherwise harmless substance.
Despite these caveats, a sizable number of species tolerate flooded and
submerged conditions well to a large extent and constitute a vigorous and
ecologically significant aquatic or amphibious flora in many parts of the World
(Maltby, 1991). Rice ( Oryza sativa L.) is the only crop plant that can easily be placed
in this group and hence its dietary preeminence in the humid tropics.
1 Department of Agricultural Sciences, University of Bristol, Long Ashton Research Station, Bristol BS18 9AF,
Great Britain
2 School of Biological Sciences, Queen Mary and Westfield College, University of London, London E14NS,
Great Britain.
ANAEROBIC GERMINATION
Rice is well-known for its ability to germinate without oxygen. In this regard
it has few rivals only since the number of other species known to germinate
anaerobically is small. These include Erithrina caffra, Nuphar luteum and Scripus
mucronatus (Menegus et al., 1992a), and various species of Echinochloa some of which
are weeds in rice field (Barrett and Seaman, 1980). In each case only the shoot emerges
from the anoxic seeds. Thus, anaerobically germinating seedling is without an
anchoring rootlet. This can be a problem in fields to direct-sown rice where anaerobic
conditions make it difficult for the crop to become firmly established. The reason
for the failure of the emerging embryonic root under anaerobic condition is not clear,
however it survives, and quickly elongates when oxygen is made available (Kordan,
1977). However, one species, Trapa natans (the water chestnut), is able to germinate
by radicle emergence (Menegus et al., 1992a). This is the only species known where
root growth is not inhibited by anoxia. On the contray in seedling, root extension is
actually promoted by the absence of oxygen in this plant. The metabolic and
physiological basis of this unique characteristic are not yet fully understood. Metabolic
studies (Menegus et al., 1992a) of T. natans have shown a high rate of alcoholic
fermentation (20 u mol ethanol g-l FW h-1 ), the availability of respirable starch reserves,
and an ability to avoid cytoplasmic acidification achieved through. (i) minimal
production of potentially fatal lactic acid, (ii) proton consumption in conversion of
malate to succinate, (iii) arginine to putrescine, (iv) glutamate to Y-aminobutyric acid
(GABA). These metabolic features are strongly reminiscent of the rice coleoptile
(Menegus et al., 1992b; Fan et al., 1993; Perata et al., 1992; Ricard et al., 1991). They are
considered to help in avoiding acidification of the cytoplasm, maintain sufficiently high
levels of ATP and dissolved solutes to support cell expansion and membrane integrity,
while other energy consuming processes such as the overall levels of protein synthesis
(Mocquot et al., 1977) and fatty acid production (Vartapetian et al., 1978) are suppressed.
However, a large number of proteins can be synthesised anaerobically in the coleoptile
(Chirkova and Hoang 1981; Atwell and ApRees, 1986), and two dimensional gels of
radio- labelled proteins synthesised in aerobic and anaerobic conditions show
remarkably similar patterns (Ricard and Pradet, 1989).
One way of improving the performance of germinating rice seeds would be
to induce the embryonic root axis to elongate as a coleoptile, while retaining the
positive gravitropism typical to aerobic rice roots. This may be possible because
anoxia and root growth are not mutually incompatible in Trapa natans. A large
32
Fig. 1.
33
Fig. 2.
34
Extension growth by the coleoptile of rice and Echinochloa oryzoides seedlings grown in air
(Air), air containing 1.0 Pa ethylene (Ethylene), or in nitrogen gas (No oxygen) for up to
8 d (taken from Pearce and Jackson, 1991).
the submergence tolerant rice cultivar 'Calrose' the length of coleoptile was 3.4 cm
longer in nitrogen environment than air after 8 day. This growth is unlikely due to
endogenous hormones such as a uxin, ethylene or gibberellins since they are inactive
when supplied to anoxic coleoptiles (Jackson and Pearce 1991; Horton, 1991).
However, the diamine putrescine appears to play a positive role in anaerobic
coleoptile elongation (Reggiani et al., 1989) while abscisic acid could be inhibitory
(Horton, 1991). Futhermore, interference in the already weak gravitropism in
anoxic
rice
coleoptile
by
auxin
transport
inhibitor
such
as
N-1
naphthylphthalamic acid (recently reported by Horton, 1994) implies that
endogenous auxin has some role under oxygen-free conditions. However, it is
contrary to competing weed E. oryzoides, where coleoptile elongation is neither
stimulated nor inhibited in absence of oxygen (Fig. 3) In these species, a 20- mm
long coleptile can easily be obtained regardless of the availability of oxygen or
the presence of elevated levels of ethylene (Fig. 3) or carbon dioxide (Pearce and
Fig. 3.
35
Condition
Coleoptile
length
(mm)
Ethylene
production
(nl g-1 h-1)
ACC
(nmol g-1
fresh wt.)
No oxygen for 4 d
29.7
00
23.1
Air for 4 d
18.7
2.1
02.5
32.0
3.7
11.7
The surrounding water then entraps the newly synthesised ethylene (Ishizawa
and Esashi, 1984), and the presence of some oxygen also allows cells to respond
36
Water
Gas phase
Aerobic
6.2*
4.5
Anaerobic
3.3
3.9
leaves
of the
since
1987)
37
Fig. 4.
38
Day
Shoot
wt(mg)
Rice
Root
wt(mg)
Seed
wt(mg)
Shoot
wt(mg)
7.4
3.2
11.8
5.0
3.7
2.4
Not Submerged
14.4
5.3
6.6
22.0
9.6
2.1
Submerged
8.9
2.6
5.6
6.9
3.0
2.3
24.7(15.1) 2.1(0)
Day 0
Echinochloa
Root
Seed
wt(mg)
wt(mg)
Day 3
Day 6
Not submerged
33.0(11)
De-submerged
13.4(4.5) 6.5(3.9)
6.2(0.6)
Plants were 7-d old at the start; Figures in parentheses are changes over the last 3 days of the
experiment.
Besides, seedlings submerged in light survive and gain a small increase in dry
weight. E. oryzoides has a superior ability to fix carbon dioxide (Smith and Walker,
1980). This may be expected from a C4 species compared to a C3 plant rice because
a C4 plant has low compensation point (Madsen, 1993). The difference between the
two species in their dependence on external carbon dioxide is shown in Fig 5. If
seedlings of both the species are grown with their shoots in moist air from which
carbon dioxide has been removed completely, growth in dry mass by the root and
39
Fig. 5.
Effect of absorbing carbon dioxide from air on growth in dry mass by initially 7-d old light-grown
seedling with the roots in nutrient solution and the shoots in a gas phase flowing at approximately
2L min -1 at 850 u mol m -2 s-1 PAR. Figures above the bars are relative growth rates (mg
mg-1 d-1 ). From Pearce, 1990.
shoot stops in E. oryzoides. But in rice, root and shoot growth continues but at the
expense of seed reserves. Thus, submerged E. oryzoides achieves its small increase
in mass by fixing carbon dioxide from the water and not by using seed reserve. In
contrast, rice while being unable to fix carbon dioxide from the dissolved water,
compensates it by drawing from the seed reserves. This way both the species survive
three days total submergence in well-illuminated water at the 7-leaf stage and are
able to grow strongly if shoots are exposed to air (Table 3).
40
CONCLUSION
Rice can germinate without oxygen, and its seedlings can grow in submerged
conditions that are partially depleted of oxygen. The coleoptile elongates to a greater
length than in normal in response to anaerobiosis, partial oxygen shortage, and
enriched ethylene or carbon dioxide. These factors are inhibitory for most of the
species. However, roots fail to emerge from rice seeds unless some oxygen is present,
while anaerobic stimulation of coleoptile elongation is limited to only a few
millimetres. Forthermore, extension of the first true leaves is dependent on oxygen,
although its effect is increased by ethylene and carbon dioxiode . If the shoots of
small seedlings are submerged, photosynthetic fixation of carbon dioxide from the
flood water is minimal and the shoot and roots depend on seed reserves for survival
over at least for 3 days.
Certain other species are shown to perform better than rice in several aspects.
Trapa natans can produce a root when germinating anaerobically and stems of
anaerobic Potamogeton pectinatus tubers elongate much more vigorously for longer
period than rice coleoptiles and have stronger negative gravitropism. The coleoptile
of Echinochloa oryzoides can extend its full length irrespective of presence or absence
of oxygen whereas the leaves of P. pectinatus can elongate strongly with oxygen. The
shoots of E. oryzoides are able to survive independently of the seed reserves when
submerged for 3 days and grow more vigorously than rice after exposed to resubmergence. Studies show how these species out-perform rice under conditions of
poor aeration, and ultimately, may help select or create strains of rice with superior
performance in flooded conditions. These studies show the capabilities of the species
other than rice to perform better under the poor condition of aeration and thus open
the scope for developing better strains of rice with such characteristics to perform
well under flooded conditions.
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Horton R F (1994) Gravitropism in rice seedlings: the effects of anoxia and N-1 naphthylphthalamic acid.
Plant Sci. 95:41- 47.
41
Ishizawa K, Esashi Y (1983) Co-operation of ethylene and auxin in the growth regulation of rice coleoptile
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Ishizawa K, Esashi Y (1984) Gaseous factors involved in the enhanced elongation of rice coleoptiles under
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Pages 47-67. In: Plant life under oxygen deprivation. Ecology, physiology and biochemistry. M.B. Jackson,
D.D. Davies and H. Lambers, eds. SPB Academic Publishing, The Hague, The Netherlands.
Jackson M B, Fenning T M, Jenkins W (1985) Aerenchyma (gas space) formation in adventitious roots of
rice ( Oryza sativa L.) is not controlled by ethylene or small partial pressures of oxygen. J. Exp. Bot. 36:
1566-1572.
Jackson M B, Waters I, Setter T, Greenway H (1987). Injury to rice plants caused by complete submergence;
a contribution of ethylene J. Exp. Bot. 37 : 1826-1838.
Justin S H F, Armstrong W (1991) Evidence for the involvement of ethylene in aerenchyma formation in
adventitious roots of rice ( Oryza sativa L.). New Phytol. 118: 49-62.
Konings H, Jackson M B (1979) A relationship between rates of ethylene production by roots and the
promoting or inhibiting effects of ethylene on root elongation. Z Pflanzenphysiol. 92: 385-397.
Kordan H A (1974) Patterns of shoot and root growth in rice seedlings germinating in oxygen. J.Plant
Ecol. 11: 685- 690.
Kordan HA (1975) Relationship between oxygen availability and transverse and vertical shoot geotropism
during germination of submerged rice seedlings. Ann. Bot. 39: 249-256.
Kordan H A (1976) Mitotic activity in rice seedling under oxygen deficiency. J. Cell Sci. 20 57-59.
Kordan H A (1977) Latent effect of anaerobiosis on root growth in germinating rice seedlings. Plant Sci.
Lett. 9 53-56.
Kordan H A, Ashraf M (1990) Environmental anoxia is unnecessary for inhibiting chloroplast
photomorphogenesis in rice coleoptiles ( Oryza sativa L.). J. Exp. Bot. 41 : 435-440.
Ku H S, Suge H, Rappaport L, Pratt H K (1970) Stimulation of rice coleoptile growth by ethylene. Planta
90 : 333-339.
Kutschera U, Siebert C, MasudaY, Sievers A (1990) Effects of submergence on development and
gravitropism in the coleoptile of Oryza sativa L. Planta 183: 112-119.
Madsen T V (1993) Inorganic carbon assimilation and growth of aquatic macrophytes. Pages 267-285. In:
Interacting stresses on plants in a changing climate. M.D. Jackson and C.R. Black, eds. Springer-Verlag,
Berlin, Germany
Maltby E (1991) Wetlands-their status and role in the biosphere. Pages 3-21 In: Plant life under oxygen
deprivation. Ecology, physiology and biochemistry M.R. Jackson, D.D. Davies and H. Lambers, eds. SPB
Academic Publishing, The Hague, The Netherlands.
Mapelli S, Bertani A (1993) Endogenous phytohormones and germination of rice under anoxia: indoleacetic
acid and abscisic acid. Pages 353-363. In: Interacting stresses on plants in a changing climate. M.B. Jackson,
C.R. Black, eds. Springer-Verlag, Berlin, Germany
Mayne G R, Kende H (1986) Glucose metabolism in anaerobic rice seedlings. Plant Sci. 45:31-36.
Mehlhorn H (1990) Ethylene-promoted ascorbate peroxidase activity protects plants against hydrogen
peroxide, ozone and paraquat. Plant Cell Environ. 13:971-976.
Menegus F, Cattaruzza L, Scaglioni, Ragg E (1992a) Effects of oxygen level on metabolism and development
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and low tolerance to anoxia. Pages 53-66. In: Surviving hypoxia: Mechanisms of control and adaptation.
P.W. Hochachka, ed. CRC Press, Boca Raton, USA.
42
Mocquot B, Pradet A, Litvak S (1977) DNA synthesis and anoxia in rice coleoptiles. Plant Sci. Lett. 9 :
365-371.
Opik H (1973) Effect of anaerobiosis on respiratory rate, cytochrome oxidase activity and mitochondrial
structures in coleoptiles of rice (Oryza sativa. L.). J.Cell Sci. 12 : 725-739.
Pearce D M E (1990) A comparison of the responses of seedlings of rice and Echinochloa oryzoides to
submergence. PhD dissertation, University of Bristol, Great Britain.
Pearce D M E, Jackson M B (1991) Comparison of growth responses of barnyard grass ( Echinochloa oryzoides)
and rice (Oryza sativa ) to submergence, carbon dioxide and oxygen shortage. Ann. Bot. 68:201-209.
Pearce D ME, Hall KC, Jackson M B (1992) The effect of oxygen, carbon dioxide and ethylene on ethylene
biosynthesis in relation to shoot extension in seedlings of rice ( Oryza sativa) and barnyard grass ( Echinochloa
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43
India
Observations
*
Growth and Yield : Tillers, biological and grain yields, grain sterility etc.
Flooding/submergence
*
The performance of some of the elite rice lines was assessed under
prolonged partial submergence during 1992 and 1993 wet seasons under
field conditions.
Observations
*
relative
Salinity/sodicity
The experiments were conducted in two situations: the normal soil which was
moderately alkaline (pH 8.5) and partially reclaimed sodic soil which had a pH 9.7.
Rainfed lowland rice genotypes were planted in field during 1992 (64 lines) and 1993
(55 lines). The details of experiments are given below :
*
Observations
*
in many parts of rainfed rice growing areas arc uncertain, which cause mild to severe
water deficit during different phases of plant growth. About half of the 38 million
ha of the rainfed lowland rice ecosystem in the world has been classified either as
drought prone or drought and submergence prone (Garrity et al., 1986).
A number of soil parameters which affect plant establishment and growth
under drought prone rainfed lowland areas need to be measured. These include soil
matric potential, hydraulic conductivity, bulk density, field capacity, wilting point
etc.
Water availability and uptake by rice plants decrease as soil matric potential drops
below zero i.e. the matric potential of saturated soil. A careful monitoring of soil matric
potential and leaf water potential is required because ample reduction in leaf expansion,
tillering and leaf photosynthesis occurs long before leaf rolling and wilting, the two
common symptoms of drought stress. CO2 assimilation is Educed by soil water potential
of only -0.5 bar (OToole and Baldia, 7982). Generally, impact of water deficit was not
observed until the soil matric potential dropped below -1 bar and leaf water potential
below -1.5 bar when leaf rolling generally begins (Setter et al., 1993).
Table 1.
(Abbreviations : S-single measurement; 2W-twice weekly; W- weekly; 2devery 2 days. d- daily; E-essential assuming equipments are available, D- desirable).
Measurements
2W
Frequency
W
2d
Stress
Drought Flooding
Salinity
1- SITE DESCRIPTION
a.
b. Topographical location
including slope
2- SOILS
a.
X
X
Continued ................
47
Measurements
S
2W
Frequency
W
2d
Drought
f.
Stress
Flooding
D
Salinity
E
g. Hydraulic conductivity/
percolation rate
3- FLOOD WATER
a. Flood water depth-rate
of increase or decrease
c.
e.
f.
EC (including source of
flooding)
g.
O2 concentration (monitor at
water and soil surface and half
way within canopy; before
dawn and at mid afternoon)
h.
CO2 concentration(monitor
as per O2)
i.
Ethylene concentration
(monitor as per O2 )
Rainfall by experiment
E
E
b.
c.
Radiation
d.
e.
f.
Evaporation by site
Continued................
48
Measurements
2W
Frequency
W
2d
5-
a.
Percent survival
d. Yield components
Stress
Drought Flooding
Salinity
f.
X
X
E
E
E
E
i.
j.
E
D
49
Fig. 1.
50
Effect of water deficits on yield in rice genotypes (per cent reduction over unstressed at top
of the bars.)
Fig. 2.
Total grains and per cent sterility (at the top of bars) in rice genotypes under water deficits.
Stress at tillering induced irrepairable loss in total biomass of all the genotypes,
but the grain yield was reduced more due to water deficit at booting stage. A
combined stress was more detrimental, but the effect was less severe in the tolerant
genotype, Kachani (Figures 1 and 2) and it was possibly due to appearance of PWP
at still lower water potential. Grain sterility was the main component responsible
for yield reduction which was 61% in IR 28 against only 32% in Kachani with stress
at tillering + boot stages. In order to quantify the stress response and explore the
mechanisms of tolerance/susceptibility to drought, the environmental parameters
listed in Table 1 need to be measured.
Effect of drought varies with the genotype, the phasic development at which
plants experience stress, the severity and the duration of drought. Rice yields are
most susceptible to water deficits at flowering than at vegetative phase (Ram et al.,
1988; Hsiao, 1982; Reyniers et al., 1982). Drought at vegetative stage causes
irrepairable loss of canopy whereas at flowering stage, water deficit hampers anthesis
and seed setting leading to higher spikelet sterility and lower yields (Ram et al.,
1988). Drought at flowering also reduced effective leaf area and photosynthesis, thus
plants have to depend on pre anthesis reserves which may impart tolerance against
internal water deficits (Chaturvedi and Ingram, 1988; Austin et al., 1980).
Furthermore, if stress coincides with anthesis, the panicle excertion may be inhibited,
spikelet sterility occurs (Ekanayake et al., 1989; Ram et al., 1988) and yields decrease
without any scope for recovery. Data on panicle emergence, and spikelet sterility,
thus provide an indication of timing and intensity of drought (Setter et al., 1993).
Drought is often associated with high temperature and intense solar radiation
alongwith low relative humidity during the dry spells in the rainy season. Soil cracks
develop and strength and bulk density increase, restricting the availability and
uptake of water and nutrients by the plants. Sizeable reductions in soil water
potential due to drought may increase salt concentrations in the soil solution which
may induce dehydration of tissues leading to cell collapse and death of plants.
51
Table 2.
Soil moisture content (% of dry weight) and leaf water potential (-bar)
in rice genotypes under water deficit condition (Pot expt.)
Stages of stress:
IR-28
(Susceptible)
Genotypes
IRAT-142
(Moderate)
KACHANI
(Tolerant)
LSD (P 0.05)
6.8
19.0
8.9
6.4
20.9
8.3
6.6
21.2
8.1
N.S.
0.60
N.S.
7.9
20.0
9.9
7.4
21.0
9.4
7.9
21.5
9.3
N.S.
0.58
N.S.
7.9
21.5
9.9
7.4
22.8
9.0
7.8
23.0
9.0
N.S.
0.60
N.S.
28.31.3
11.20.9
28.351.1
28.31.3
10.10.95
28.01.2
28.11.0
9.80.72
27.50.95
Soil moisture
At field capacity
At PWP
After revival
Weed populations under drought invariably exert a threat to crop stand and
render plant easily accessible to insects and diseases. Hence, data on temperature,
relative humidity and light intensity above and inside the plant canopy, weed, insect
and disease profiles may also be included in the environmental measurement data
set (Table 1). All these environmental variables can effectively be integrated through
simulation models for understanding the response of drought on crop growth and
productivity. Such simulation models have been suggested by a number of workers
which include measurements on soil, climate and plant factors (Penning de Vries et
al., 1989; Woopereis et al., 1993; Singh, 1993).
With the above set of environmental and plant measurements, one can easily
predict the timing, duration and intensity of water deficits. This knowledge may
enable breeders to synthesize a new rice genotype suitable for drought prone rainfed
lowlands.
Flooding
Flooding is a serious constraint to rice yields in rainfed lowland and deepwater
areas due to partial or complete submergence of plants. Partial submergence is
defined as when 40 to 99 per cent shoot is submerged in water. Flooding in rice
fields is of two types : i) Flash or intermittent (ii) Stagnant or prolonged.
Approximately 22 m ha of rice is affected by flooding which includes 15 m ha of
52
flash flood areas of rainfed lowland rice and 5 m ha of deepwater rice (Khush, 1984).
In India, approximately 60% of Kharif rice area is subjected to waterlogging due to
flash and stagnant floods (Balkrishna Rao and Biswas, 1979).
Flash flooding refers to a situation where the rice crop gets submerged due
to sudden increase in floodwater for varying periods, normally not exceeding 10-12
days (Dwivedi and Hille Ris Lambers, 1991; Senadhira, 1992). The period of flooding
is often prolonged in Eastern India. Flash flooding normally occurs near river,
riverlets and streams. The rice area near Ghaghara, Ganges, Rapti, Gandak, Kosi,
Baiterni, Mahanadi, Rishikulya, Brahmaputra, Krishna and Cauvery rivers and in
the tidal wetlands coastal area of India, are more vulnerable to flash flooding (Singh,
1982). The flash flooding occurs mainly during vegetative stage of rice and its severity
depends on the intensity and duration of rainfall, nature of water source and
topography.
Rice, by nature, is tolerant to waterlogging, flash flood and associated anoxia
which makes its cultivation possible under flood prone lowland ecosystem. A
number of morphological, physiological and biochemical features are responsible
for tolerance to submergence which are influenced by genotype, submergence depth
and duration and other environmental conditions. Features, like coleoptile, that
tolerate anoxia (Taylor, 1942) and the ability of the coleoptile (Ohwaki, 1967) and
stem (Vergara et al., 1976) to elongate vigorously when submerged, provide chance
to escape complete inundation or minimize the duration. Other attributes include a
root system that can elongate more rapidly in the presence of ethylene (Konings and
Jackson, 1979), a highly porous morphology that facilitates internal diffusion of
oxygen down the plant (Armstrong, 1971) and a hydrophobic outer surface that
traps a coating of air when leaves are submerged. This forms a gaseous film that
may facilitate basipetal diffusion and mass flow of oxygen from the aerial
environment when some leaf tissues remain above the water surface (Raskin and
Kende, 1984).
Despite these adaptive features, most rice cultivars can not survive complete
submergence for a long time, more so, during early stage of development. The
experiments conducted at NDUAT and elsewhere indicate that 45-day old seedlings
showed better tolerance to submergence (7, 11 and 15 days submergence) than 30
or 15-day old seedlings with good genetic variability (Mazaredo and Vergara, 1982;
Chaturvedi et al., 1993).
Experiments conducted at the consortium site at Masodha indicated that with
similar experimental conditions and crop management, rice yields among genotypes
during 1992 and 1993 wet seasons showed a variation of 0.2 to 2.9 t/ha (Table 3). It
is clear from the data that environmental factors like water depth and duration of
submergence, temperature, light, rainfall and humidity have an important role in
changing the phenology, for example, flowering delayed during 1993. (Figures 3,
and 4). Environmental characterization (Table 1) thus, is of utmost importance for
interpreting the performance and determining the stability of the genotypes across
locations and at the same location over different years.
53
Table 3.
Performance of elite
submerged condition.
Genotypes
lines
50% flowering-DAS*
of rainfed
lowland
ERT*/hill
rice
under partially
Yield t/ha
Yield
Difference
1992
1993
1992
1993
1992
1993
7.5
5.3
7.2
+1.9
NDR-30030
113
122
7.8
NDR-30076
118
121
8.2
7.4
5.2
7.0
+1.8
NDR-30039
115
118
8.4
8.1
4.5
6.0
+1.5
NDR-30023
112
118
9.3
7.8
4.3
6.2
+1.9
NDR-40013
120
127
6.7
6.7
4.8
6.4
+1.6
NDR-40032
114
123
8.4
7.4
4.5
7.4
+2.9
Madhukar (Check)
115
126
5.2
7.0
2.0
1.8
-0.2
SEm
0.66
0.71
0.24
0.14
0.23
0.15
LSD (P0.05)
207
223
0.75
0.43
0.72
0.47
**
Earbearing tillers
There were large differences in water depth (Figures 3 and 4), showing lower
water levels (mean depth 5.9-11.0 cm) during July and August in 1992 (active tillering
phase) and deeper water level in 1993 (mean depth 19.8 to 23.9 cm). In contrast,
during 1992 higher water depths (11.0-34.5 cm) were observed at flowering to milk
stages than in 1993 (4.0-26.5 cm). The physiological implication of such variations
needs to be looked into as a key to the effect of environmental factors affecting
growth and yield. Water temperature variations and sunshine hours should also be
taken into consideration.
A number of physiological and biochemical explanations for death and
survival under submergence has been reported which include a possible interaction
between CO2 low O2 and high ethylene concentrations (Setter et al., 1988, 1989).
Low photosynthesis due to low CO2 diffusion leads to lower carbohydrate
assimilation and energy deficiency which restrict nutrient uptake and growth.
Submergence may also result in solute leakage due to low O2 concentration and
affect membrane permeability (Smith and ap Rees, 1979).
The rice plants respond and adapt to various types, timing and duration
of flooding in different ways which include elongation for deepwater rice
varieties and submergence tolerance for lowland varieties exposed to transient
flash floods (Hille RisLambers and Seshu, 1982).
A number of environmental and plant factors are associated with damage due
to flooding namely low light, siltation on the leaves, mechanical damage, solute
leakage, limitation to gas diffusion, accumulation of toxic metabolites inside the
plant, insect-pests and diseases (Creenway and Setter, 1994). Additional adverse
54
Fig. 3.
Variation in daily water depth during 1992-93 wet season crop at C.R.S. Masodha, India.
55
Fig. 4. : Weekly meteorological information during wet crop season (1992 & 1993) at C.R.S. Masodha, India.
effects may be associated with desubmergence and post hypoxic injury (see
Crawford, 1992, for review). These factors seldom act alone since alteration in one
component may lead to a number of other adverse effects which ultimately affects
survival and growth of plants under submerged conditions.
Submergence at seedling stage is most harmful (reducing plant population),
followed by submergence at tillering stage which cuts down the tiller production.
Submergence at boot and flowering stages prevents panicle emergence and increases
spikelet sterility (Pandey et al., 1979). High nitrogen level in the soil accentuates
adverse effects of submergence (Palada and Vergara, 1972). Gas diffusion during
submergence is quite low in water, hence any gas (CO2 ethylene and methane)
which is produced under water increases in concentration, while that O2 consumed
decreases in concentration. Any of these changes may reduce growth, and survival
of rice during flooding.
The composition of floodwater may vary with locations, hence proper
measurement and characterization of flood water is essential in order to assess its
effect on plant growth and yield. During 1993 wet season, a brief environmental
characterization of stagnant water in submerged rice fields was done at crop research
stations, Pagalabhari, (Faizabad) and Ghagahraghat, (Bahraich) (Figures 5, 6 and 7).
A diurnal variation in O 2 concentration of water at different depths was noted
showing a decrease in O2 with increase in water depth (0.01 - 0.4 m). Maximum O 2
in water was observed during mid day possibly due to photosynthesis of algae and
submerged rice plant tissues which gradually declined to the lowest before dawn.
Plant respiration during night might have consumed at least some of the dissolved
oxygen (Figure 5). In deepwater, a gradual decline in dissolved O2 was noted upto
1.4 m depth, beyond which, a sharp decrease occurred leading to zero oxygen at 2
m depth well above the soil surface (Figure 6). The temperature profile through
water depth at 10.00 h during day decreased with depth but it ranged only between
27.5 to 28.0 C in the deepwater at Ghagharaghat.
Fig. 5.
57
Fig. 6.
Oxygen concentration of flood water in deep-water rice field at Ghagharaghat, Bahraich, India.
Maximum water depth was 2.4 m.
Fig. 7.
58
Salinity
An estimated 150 m ha of current and potential rice lands in the tropics and
subtropics are affected by salinity (Massoud, 1974). in South and South East Asia
alone about 90 m ha of lands suited to rice production, lie idle largely because
of soil toxicities of which nearly 49 m ha are saline and 12 m ha are alkali
(sodic) in nature (Ponnamperuma and Bandyopadhya, 1980). In general, soil
salinity is caused by presence or by intrusion of sea water or by surface
evaporation of brackish soil water. Salt accumulation increases in the drier
climates and diminishes strongly in equatorial climates without a pronounced
dry season.
Rice is generally reported to be a moderately salt tolerant crop but no rice
variety can withstand high salinity or sodicity throughout its growth cycle. High
content of sodium chloride in soil solution with specific conductance values of
more than 8-10 dS/m are harmful to rice plants and cause as much as 50% yield
59
reduction. In tidal areas, the specific conductance values change from day to day
depending on tidal regimes (De Datta, 1981).
Inland saline sodic soils generally have pH more than 7 alongwith high
amounts of soluble salts. The underground water invariably is brackish. Saline-sodic
soils have high amounts of soluble salts along with high concentrations of carbonate
and bicarbonates of sodium. High sodium replaces calcium from the clay complex
with concurrent precipitation of calcium carbonate which forms an impermeable
(upto 1 m thick) kankar layer. The soil becomes strongly dispersed and hence highly
impermeable to water. This necessitates the careful characterization of coastal and
inland saline and sodic soils in terms of topography, bulk density, hydraulic
conductivity, variations in pH, electrical conductivity (ECe) and chemical properties
at essential and desirable levels. Salt affected soils show a number of problems which
hamper normal growth and development of plants (Ponnamperuma and
Bandyopadhya, 1980).
It is evident from the data presented in Tables 4 and 5 that a large variability
exists in the physico-chemical properties of soil including nutritional status which
should be accurately characterized if the results obtained at one location are likely
to be extrapolated to other locations. Underground water is often brackish at certain
locations and this too needs careful monitoring.
Table 4.
Soil parameter
0-15
15-30
Sodic
Normal*
9.5
8.5
9.8
8.7
ECe (dS/m)
Sodic
Normal*
1.4
0.6
1.8
1.0
Sodic
Normal*
35
15
55
17
Sodic
Normal*
0.8
2.5
0.3
2.0
Sodic
Normal*
14-16
16-19
13-15
14-17
Sodic
Normal*
silty loam
silty loam
Topography
undulating
60
Normal designates a soil where 10 t/ha pyrite was applied to sodic soil two years earlier,
followed by continuous cropping.
Soil parameters
1530
0.050.2
0.20.3
Sodic
Normal
0.10.2
0.30.4
Sodic
Normal
20
25
AvailableP2O 5 (ppm)
Sodic
Normal
28
25
Sodic
Normal
300
320
Iron
Sodic
Normal
3.8
9.5
3.7
7.2
Manganese
Sodic
Normal
2.9
3.0
2.3
2.7
Zinc
Sodic
Normal
2.2
5.6
1.2
4.5
CaCO3 (%)
Sodic
Normal
6.0
1.5
5.5
2.0
Thickness of CaCO3
(Kankar) layer
Sodic
present
(2540 cm thick)
rarely present
present
Normal
good
greatly depending
to survival and
Eduction in grain
A large variability
61
was also noted in dry matter partitioning into grains indicating a reduced CO2
fixation or reduced translocation of carbohydrates under high salt concentrations.
Tolerant genotypes had lower salt injury index (0.11 - 0.29) against 0.48 - 0.71 in
susceptible genotypes (Table 7). A comparison of yield versus trait combinations
(Table 8) and harvest index versus trait combinations (Table 9) indicates that
interaction between biological yield, harvest index and grain fertility plays an
important role in determining yield of the genotype. Genotypes with higher total
biomass and better partitioning may be looked into as the future cultivars for salt
affected rice areas.
Table 6.
Genotype
% reduction
over normal
Normal Soil
(PH 8.5)
Sodic Soil
(PH 9.7)
Usar-1
73.0
70.2
3.8
CSR-10
56.2
49.7
11.6
TCA88-10-1
70.0
54.1
22.7
NDR-80
66.0
61.5
6.8
Purple
58.0
46.3
20.2
NDRK-5003
94.0
65.7
30.1
Pokkali
63.7
43.3
32.0
Pusa- 516
79.0
42.0
46.8
Rajshree
64.0
33.4
47.8
Kasturi
67.0
28.1
LSD (P0.05)
9.77
6.82
58.1
-
Tolerant
Susceptible
The literature frequently contrasts the central dilemma of ion toxicity versus
water deficit (osmotic effect) in saline conditions (Flowers et al., 1977 for halophytes
and Greenway and Munns, 1980 for non-halophytes). Large quantities of salts are
carried through the transpiration stream to the leaves which eventually leads to their
death. NaCl accumulation in leaves is correlated with reduced photosynthetic
activity and with ultrastructural and metabolic damage (Yeo and Flowers, 1986,1989)
and there is now direct evidence that this is mediated by ayoplastic increase in salt
62
concentrations in the expanded leaves (Flowers et al., 1991). There is ample variability
in salt susceptibility between (Yeo et al., 1990) and within (Yeo el al., 1988) varieties,
differences in vigour accounting for much of the variation in the survival in salinity
(Yeo et al., 1990). Short term initial effects of salinity are primarily due to limitation
of water supply, whereas long term effects are due to accumulation of salt within
expanded leaves (Yeo et al., 1991).
Table 7.
Grain Yield and salt injury index in tolerant and susceptible rice
genotypes.
Genotype
Harvest
index (%)
at pH 9.7
Salt injury
index
Normal Soil
(pH 8.5)
Sodic Soil
(pH 9.7)
Usar-1
24.7
22.0
31
0.11
CSR-10
26.0
20.7
44
0.20
CSR-11
26.9
22.2
33
0.17
TCA 88-10-1
24.3
20.0
37
0.17
NDR-80
26.1
22.6
37
0.13
Purple
24.7
21.9
43
0.11
NDRK-5003
24.5
19.9
30
0.19
Pokkali
22.1
15.5
34
0.29
Pusa-516
23.5
12.2
23
0.48
Tolerant
Susceptible
Rajshree
34.2
10.1
30
0.71
Kasturi
21.1
8.4
30
0.60
LSD (P0.05)
3.96
3.00
4.55
presented in Table-10 indicate the role of beneficial ions like K+ and Ca2+ in mitigating
the detrimental effects of high internal Na + . Almost all the salt tolerant genotypes
invariably possessed higher K/Na and Ca/Na ratios than susceptible ones. A
number of mechanisms for exclusion and sequestering/compartmentation of ions
for mitigating the adverse effects of salinity/sodicity have been proposed by Flowers
el al. (1977, 1991) and Yeo and Flowers (1984) including exclusion, sequestering of
ions in roots, old leaves, or different cellular compartments or dilution by growth.
Table 8.
Genotype
Biol. Yield
(g/hill)
HI (%)
Grain
fertility (%)
EBT/hill
Salt
injury
index
NDR-80
22.6
61.5
37
87
12
0.13
Usar-1
22.0
70.1
31
90
16
0.11
CSR-11
22.2
56.5
33
83
15
0.17
Purple
21.9
46.3
43
85
18
0.11
CSR-10
21.7
49.7
44
74
16
0.16
CSR-18
21.8
50.0
44
90
15
0.19
IET-12856
22.8
55.1
40
90
12
0.17
IET-12865
21.0
53.0
39
82
12
LSD (P0.05)
N.S.
3.10
4.66
8.15
4.95
0.19
-
Table 9.
Genotype
HI (%)
Yield
(g/hill)
Biol Yield
(g/hill)
Grain
fertility (%)
EBT/hill
Salt
injury
index
Usar-1
31
22.0
70.1
90
16
0.11
NDRK-5003
30
19.9
65.7
88
12
0.18
NDRK-5002
30
17.1
56.5
79
11
0.37
Rajshree
30
10.1
28.1
46
11
0.70
Kasturi
30
8.4
38.2
43
10
0.60
IET-13408
30
15.9
54.0
91
10
0.47
IET-11353
30
14.3
47.2
80
11
0.25
N.S.
3.68
2.85
6.67
2.71
LSD (P0.05)
64
Table 10.
Genotype
K/Na and Ca/Na ratios in salt tolerant and susceptible rainfed lowland
rice genotypes.
Leaf
K/Na ratio
Stem
Root
Leaf
Ca / Na ratio
Stem
Root
Tolerant
Purple
2.4
1.9
1.1
3.0
4.1
2.1
CSR-10
3.0
3.0
3.1
2.0
2.5
2.2
CSR-5
2.5
3.1
2.5
2.1
2.5
2.2
CSR-18
0.8
1.6
2.1
2.6
Pokkali
1.5
1.7
1.7
1.3
1.8
1.7
TCA-88-10-1
1.9
1.6
1.5
1.3
2.8
1.7
IET-11351
16
1.2
1.2
2.5
1.8
1.6
Sajoo-52
1.6
1.4
2.9
2.2
Pusa-516
1.1
0.9
0.7
0.9
1.0
0.9
Basmati-370
1.0
0.8
1.3
1.1
IET-13407
0.9
0.7
0.9
1.4
1.4
1.1
Susceptible
A distinction should be made between saline soils which contain enough Ca2+
to meet the nutritional requirements of the plants and sodic soils, in which very little
available Ca2+ is present while the concentration of exchangeable sodium is greater
than 15% and ample carbonate and bicarbonate ions are present leading to high pH
of the soil solution. In salt affected soils, accumulation of Na+ occurs in the stem
and leaves of the plants which vary with plant species. Rice accumulated more Na
and Ca than barley or wheat (George, 1967).
We have also observed a selective uptake and accumulation of Na+ , K+ and
Ca+ by different rice genotypes. Tolerant genotypes usually showed higher K/Na
and Ca/Na ratios in leaf, stem and roots than susceptible genotypes. These
measurements may be used as selection criteria for screening rice genotypes for salt
tolerance and should be included for environmental characterization as plant
measurements (Table 1). Importance of vigour in salt tolerance of rice was
emphasized which reduced actual ion concentration due to dilution and thus
decreasing the adverse effects of salinity. Vigour parameters included for
measurement are given in Table 1.
The mechanism of injury or tolerance to coastal and inland salinities may vary,
as in the former case, the plants are exposed to intermittent high levels of salinity
due to tidal inundations which often fluctuates. Inland salinity, though also fluctuates
with other environmental factors like rainfall, temperature, humidity and light, but
plants are exposed to a continuous stress, which enables them to adjust against
slowly increasing ion concentrations within tissues. In both cases the slow build up
65
of ions to inhibitory concentrations may explain why growth reductions over short
exposure periods of salinity are less meaningful than growth or survival measured
over several weeks (Yeo and Flowers, 1984; Akita and Cabuslay, 1986).
In rice growing areas, the inland saline soils especially in India and Pakistan
are notorious due to associated alkalinity problems alongwith nutritional
imbalances. Rice plants grown in saline-alkali or alkali soils are exposed to high pH,
low to very low hydraulic conductivity, deficiency of nitrogen, zinc and iron, besides
high soluble salts. These factors create an array of complex situations for plants
growing therein. Neue et al. (1990), found that salinity tolerance scores for rice are
significantly correlated to other stresses such as alkalinity and peatiness, P deficiency
to alkalinity and zinc deficiency and Zn deficiency to Fe toxicity. Pokkali, one of
the tolerant rice varieties to salinity could not match well to CSR-10, NDR-501 and
TCA-88-10-1 in our rice screening programme for rainfed lowland salinity/alkalinity
(Tables 6, 7). The reason is clear on the basis of the soil characterization since most
of the saline soils are also highly alkaline (pH upto 10.5) and deficient in nitrogen,
zinc and iron. This result contradicts the generalization that more vigorous and tall
genotypes are tolerant to salinity (see also Tables 6 and 7). Dilution effect may be
true over short term and needs to be evaluated further in long term experiments.
The complexities of saline environments, thus, render it difficult to
characterize. The EC of the soil solution is the most important component to quantify
the overall ions, and needs a careful monitoring during the experiment. The soil
matric potential, water content, and nutrient status may also be given careful
consideration. Soil pH, hydraulic conductivity, and the nature of CaCO3 hardpan
below root zone may essentially be measured in order to seggregate the plant
responses to saline versus alkaline environments.
CONCLUSION
The forgoing discussions clearly indicate the importance of environmental
characterization for drought, flood and salinity situations having enormous amount
of variability across locations and seasons. Such measurements enable extrapolation
of findings from one site to the other sites or locations in the rice growing countries.
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69
SUMMARY
Uptake and transport of nutrient ions by roots of higher plants is strongly
dependent on oxidative phosphorylation in order to supply adequate ATP. When
aerobic respiration is slowed down by O2- deficiency in the rooting zone, roots
become hypoxic or anoxic; mineral nutrition may be affected. In barley and wheat,
phloem-mobile nutrients are reallocated from older, senescing leaves to the younger
and growing ones; but the mineral nutrition of the plant as a whole is not sufficient
and deficiency symptoms appear. Under saline conditions, Na+ fails to be excluded
by roots of maize and reaches leaves in injurious amounts. Uptake of Cl- from low
concentration, a thermodynamically active process is strongly inhibited by anoxia,
even when roots are not totally depleted of ATP.
When cereals such as barley, wheat or maize develop aerenchymatous roots
in response to O2 deficiency, nutrient uptake can continue despite the loss of
many cells in the root cortex. Internal transport of O2 from the leaves improves
the energy status of the root. In rice, and other aquatic plants that form an
extensive aerenchyma, suberization and lignification of the root axes assists in
O2 conservation by minimizing radial leakage and loss to the rhizosphere.
However, these structures sometimes act as barriers to radial, inward, transport
of nutrient ions in other species, so that uptake probably depends largely on the
formation of fine laterals that lack structural barriers.
INTRODUCTION
Most cereal crops are not well-adapted to wetland conditions, showing a
variety of responses to a sudden excess of water in the rooting zone, including
inhibition of leaf extension and tillering, precocious senescence of older leaves,
stomatal closure and reduction of net photosynthesis, and finally in a drop in
economic yield (Van't Woudt and Hagan 1957; Cannell and Jackson, 1981; Meyer et
al., 1985; Hodgson et al., 1990). Such flooding symptoms are undoubtedly a
consequence of the inadequate performance of the roots. In transiently flooded soil,
dissolved O2 is quickly depleted by the rapid respiration of roots and
micro-oganisms when temperatures are high, (Meyer et al., 1985; Hodgson et al., 1990)
1 Department of Horticultural Sciences, Texas A & M University, College Station, Texas 77843-2133, USA.
soil environment account for this? We investigated this question with barley (Drew
and Sisworo, 1979) and wheat (Trought and Drew 1980a) grown in 7.5 cm diameter
cylinders of a sandy loam soil (2% organic matter). After barley grew for 13 days
(18C day 16 C night) in a controlled environment the soil was water saturated and
the changes in the composition of the soil solution as well as various plant
parameters, were measured with time. The most significant change in the soil was
the loss of dissolved O2 , which was essentially depleted in 24 h period. The initial
symptoms of flooding damage to barley shoots, such as slow increase in fresh weight,
inhibition of leaf extension, and chlorosis of the tops of the oldest (lower) leaves
appeared in 2-4 days. After only 2 days, there was a marked decrease in concentration
of N, P and K in shoots. At that stage, none of the potential toxins that accumulate,
as the soil redox declines, had reached appreciable concentrations and there was still
ample free nitrate. It was concluded that the early responses to flooding, including
the lowering of mineral nutrient status, was the result at least in part of the inhibition
of ion transport by roots through lack of O2. The premature chlorosis or senescence
of older leaves with flooding was accompanied by a rapid loss of N from those
leaves, indicating a remobilization from older to younger leaves. With wheat in soil
columns (Trought and Drew 1980b), flooding strongly inhibited net uptake of N, P
and K into the shoots. Remobilization from older to younger leaves was found to
include all three of these phloem - mobile nutrients.
Increased concentrations of Fe and Mn in shoots in the longer term accord
with the much greater solubility in the soil water at low redox potentials, while Na +
exclusion by roots fails under O2 -deficient conditions. If O2 -deficiency itself accounts
for the early stages in flooding response in cereals, it seemed reasonable to expect
that plants in de-oxygenated nutrient solution would respond similarly to those in
flooded soil. Wheat was grown in nutrient solution for 13 days at 14C in a controlled
environment room, and then the mots were made suddenly O2 deficient by sparging
with N2 gas (Trought and Drew, 1980c). Essentially all of the responses recorded
with soil flooding occurred with plants in deoxygenated nutrient solution, where
none of the potentially harmful products of low soil redox could accumulate.
Net uptake of nutrients into the shoot was almost eliminated with
deoxygenation of the nutrient solution. The average concentration of nutrients in
the xylem sap was calculated from measurements of the volume of water transpired
and the net transfer of nutrients to leaves; for P, K, Ca and Mg these concentrations
were equal to, or less than, those in the nutrient solution, indicating that mass flow
of water through the roots to leaves could account for the small continuing transfer
of nutrients. For N, the calculated concentration in the xylem sap exceeded that of
NO3 in the nutrient solution. It may be suggested that N might be remobilized from
within the senescing root system, or be absorbed by the small number of
aerenchymatous roots that were beginning to form at the shoot base and enter the
nutrient solution toward the end of the experimental period.
If mineral deficiency of shoots is a contributory factor in flooding damage,
might it be possible to overcome some of the symptoms by delivering additional
nutrients to the shoots? It has long been recognized that under outdoor conditions,
72
heavy dressings of N to the soil surface can help alleviate the detrimental effects of
soil flooding (Van Hoorn, 1958; Belford, 1981). There is a variety of explanations for
this, including the delaying effect of NO3- on the lowering of the soil redox potential
(Ponnamperuma, 1972) and replacement of NO -3 lost in denitrification (Gambrell
and Patrick, 1978). However, with wheat we used a split root technique to show that
as long as a small part of the root system was kept oxygenated and supplied with
a full complement of nutrients, nutrient uptake and growth were comparable to that
of aerobic control plants (Trought and Drew, 1981). A remarkable alleviation of
flooding injury in soil columns was obtained with barley, simply by regular additions
of Ca (NO3)2 solution to the soil surface (Drew et al., 1979). Here, presumably the
hypoxic roots at the surface absorbed nutrients sufficiently rapidly to compensate for
the disfunction of the remainder in the deeper soil. It seem as if mineral nutrition
together with its interaction with the plants hormonal status can largely overcome the
deleterious effects on the shoot.
It has been suggested that NO3- can act as an alternative electron acceptor to
O2 under anoxia, and thereby, permit regeneration of NAD from NADH to maintain
glycolysis(Garcia-Novo and Crawford, 1973). In effect, a competition was envisaged
between nitrate reductase and alcohol dehydrogenase (for NADH) such that
anaerobic production of ethanol would be inhibited. However, experiments with
rice roots (Reggiani et al., 1985) failed to confirm this expectation. Moreover, with
maize roots no evidence could be found that the presence of NO -3 enhanced the
energy status under strict anoxia (Saglio et al., 1988); a more rapid provision of NAD
might be expected to accelerate glycolysis and substrate- linked phosphorylations
to yield ATP. Thus the effectiveness of 'nitrate respiration' in roots of higher plants
remains in doubt.
ANOXIA AND ION TRANSPORT BY NON-AERENCHYMATOUS ROOTS
Absorption of cations and anions across the plasma membrane of root cells
is linked directly or indirectly to the activity of H+ -translocating ATPases that pump
protons out of the cell towards the external medium (Poole 1978; Serrano 1989;
Sanders, 1990). The inward (passive) flux of protons, in co-transport with other ions
is the initial step in the absorption of nutrient ions into the plant. After radial
transport across the root, transport from xylem parenchyma into the xylem likewise
depends on the activity of a similar H+ -pump (Hanson, 1978; Clarkson et al., 1984).
Higher plant cells contain no reserves of ATP, which in metabolically active
cells like those of the root apical zone, is sufficient to maintain metabolism for less
than 60 seconds at room temperature (Roberts et al., 1985). The immediate effect of
sudden imposition of anoxia is thus to deplete cells of nucleotide triphosphates, and
thereby arrest the activity of proton pumps. Depolarization of membrane potential
is therefore an almost immediate response to the sudden imposition of anoxia
(Cheeseman and Hanson 1979; Buwalda et al., 1988), and presumably the dissipation
of the H+ gradient across the plasma membrane curtails active ion transport (Rao
and Rains, 1976; Cheeseman and Hamon, 1979; Jacoby and Rudich, 1980). In anoxic
maize roots, passive inward movement of K+ could still continue because of a-100mv
73
diffusion potential at the membrane (Cheeseman and Hanson, 1979), indicating that
a general degeneration of membrane properties did not take place in the short term.
This accords with other observations that the apical zone of maize mots, the zone
most sensitive to anoxia, retains viability for 15-18 hours when anoxically shocked
(Roberts et al., 1984; Johnson et al., 1989). However, the evidence suggests that
anaerobic generation of ATP during fermentation is insufficient to energize ion
transport in competition with all the other requirements for ATP in energy-depleted
cells.
With long term anoxia, a seemingly generalized decline in plasma membrane
properties takes place with loss of inorganic ions and organic solutes from roots of
wheat (Greenway et al., 1992). The concentrations of solutes remaining in the roots
at 70 hours of anoxia, as a percentage of the initial values, were K+5%, free amino
acids 41%, soluble sugars 7%. However, in experiments with roots that had been
aciclimated by exposure to solutions bubbled with about 6% O2, losses at 20 hours
of anoxia were measured for different mot zones. For the apical 1mm zone for those
mot tips that had lost their elongation potential (i.e. were moribund), there was little
retention of K+, amino acids and sugars. Root tips that could still grow, and older
mot zones (1-5mm and 10-20 mm) maintained similar concentrations of solutes as
the controls. There was a close association between generalized solute leakage and
loss of cellular viability of the root tips of wheat.
Subapical zones of hypoxically acclimated roots of wheat retain their ability
to transport ions after 24 h of anoxia, when re- exposed to air (Greenway et al., 1992).
Evidently in these relatively anoxia-tolerant cells, which do not leak readily like
anoxically shocked ones, the machinery for ion transport is not lost.
The effect of O2-deficiency on ion transport to the xylem has long been
a topic of interest. The hypothesis of Crafts and Broyer (1938) concerning the
mechanism for ion transfer from xylem parenchyma cells to the xylem
postulated that those cells bordering the xylem were leaky as a result of a
deficiency of O2 within the stele. Under most conditions of aeration, no such
deficiency of O2 occurs, and the 2-pump hypothesis is now widely accepted.
However, when aeration becomes less than ideal, there is evidence that the
stele in maize roots does become anaerobic (Thomson and Greenway, 1991) :
for enzymes like pyruvate decarboxylase and alcohol dehydrogenase that are
indicative of anaerobic, fermenting cells, there is induction in the stele but not
the outer, more aerobic cortex.
OXYGEN DEFICIENCY AND SALINITY
Flooding of the soil with saline water represents a special situation of economic
interest, because irrigated agriculture in arid and semi-arid climates is often
associated with poor soil drainage and excess salts (Carter, 1975). In terms of root
function, environments low in O2 allow a greater transfer of Na+ from roots to shoots.
With adequate oxygenation, Na+ is actively pumped out of the cytoplasm of root
cells to the outer solution, and also sequestered by xylem parenchyma cells (Shone
et al., 1969), thereby depleting Na+ in the ascending xylem sap. Such mechanisms
74
are dependent on ATP and break down when deprived of O2 (Drew and Lauchli,
1985; Drew et al., 1988). The additional flux of Na+ to the xylem, together with
inhibition of K+ transport, can lead to extreme changes in the ratio Na : K entering
the shoots. In one study by Drew et al. (1988) this ratio increased by a factor of 860
for O2 deficient roots when the concentration of NaCl in the external solution
increased from 1.0 to 50.0 mM. Absorption of C1 into roots is normally dependent
on ATP for active transport, and O 2-deficiency strongly inhibits uptake at low to
moderate concentrations (upto about 100mM) in maize (Ghosh and Drew,
unpublished). However, at higher concentrations we found that passive transport
was appreciable and perhaps as a result of depolarization of plasma membranes,
C1 translocation to the shoot was enhanced relative to aerobic controls. For salt
sensitive species like rice and maize (Yeo et al., 1988), root O2 -deficiency can thus
greatly exacerbate the detrimental effects of excess salts.
NUTRIENT UPTAKE BY AERENCHYMATOUS ROOTS
The internal transfer of O2 from shoots to roots within aerenchymatous cortical
tissue is an important mechanism for complete or partial avoidance of O2 deficiency
in many cereal species. In rice, the system of interconnected gas spaces or lacunae
is very well developed, as it is in many aquatics, so that roots can extend considerable
distances into anaerobic soil (Armstrong, 1979), but the demand for O2 is just as
great as in non wetland species (Armstrong and Webb, 1985). The low porosity to
O2 of the hypodermis, other than in the apical zone, ensures rice roots to avoid loss
of O 2 to the surrounding soil, but rather conserve it for respiration and for
oxygenation of the rhizosphere in the tip zone. Additionally, the extension of
prominent intercellular spaces into the apical meristem probably assists diffusion of
gaseous O2 to a group of cells with a very high respiratory requirement for O2
(Armstrong, 1979). Non-wetland species are not so well-adapted, failing to form an
impervious hypodermis, or such a prominent aerenchyma, or intercellular spaces in
the meristem.
In non-wetland cereals, aerenchyma formation in the roots requires induction by
hypoxia (Thomson et al., 1990). Such conditions enhance ethylene biosynthesis and
entrapment within root tissues, the ethylene in turn promoting lysis of cells in the cortex
to form interconnected lacunae (Drew et al., 1979; Jackson et al,, 1985a). Aerenchyma
formation in rice is mostly constitutive, failing to demonstrate a clear signaling by
ethylene (Jackson et al., 1985 b).
What is the significance of such changes in root anatomy to radial ion transport
to the xylem? In aerenchymatous roots of maize, radial transport of K + and Pi under
well oxygenated conditions was not inhibited by the degeneration of many of the
cortical cells Drew et al, 1980; Drew and Saker 1986). The remaining radial files of
intact cortical cells, and radial cell wall remnants, appeared able to transport ions
across the cortex without lowering the overall rate of transport (Drew and Fourcy,
1986). In situ, such aerenchymatous roots are sufficienctly well supplied with O 2 to
maintain energy metabolism (Drew et al., 1985), so that it seems reasonable to
suppose that ion transport can continue.
75
In rice and other species with strongly suberized and lignified hypodermal
layers (Clark and Harris, 1981), the question arises as to whether these act as barriers
to ion transport (reviewed by Drew, 1987). In the sand sedge (Carex arenaria), the
hypodermis becomes suberized within 2 to 5 cm of the root tip and becomes highly
impermeable to water and inorganic nutrients (Robards et al., 1979), so that uptake
is confined to the finer lateral roots lacking such anatomical features. In maize, the
permeability of the hypodermis depends on the humidity to which the mot has been
exposed, with greater impermeability resulting from exposure to air (Ferguson and
Clarkson; 1976; Peterson and Perumalla, 1984; Clarkson et al., 1987). In sorghum
exposed to dry soil, lignification and suberization of the hypodermis and endodermis
occur unusually close to the mot tip, and the late maturing metaxylem vessels fail
to lose their cross walls (Cruz et al., 1992). The combined effect is to make such roots
ineffective in water absorption. Relatively little work has been published on the
precise zones along rice seminal and nodal roots that contribute to ion transport, so
that the consequences of anatomical changes are less well understood. Such work
is best done with radio-labeled tracers (Peterson, 1987), because although apoplastic
fluorescent dyes provide useful evidence concerning changes in the resistance of
potential pathways for flow of solutes and water, their larger molecular size is likely
to become excluded where some passage of smaller molecules can continue.
CONCLUSION
For non-wetland species, oxygenation of the root system can greatly modify
its function in plant mineral nutrition. Simultaneously, changes in permeability to
water, and the synthesis of phytohormones and their transfer to the shoot, contribute
to the symptoms of flooding damage. The ability of root cells to maintain viability
under anoxia, together with the rapid formation of a new, aerenchymatous root
system of a size that can replace the function of the original, damaged one, are
features that assist greatly a plant's tolerance of transient flooding. Better
understanding of the genetic mechanisms of anoxia-tolerance and anoxia-avoidance
will help in further improvement of performance of flood-sensitive dryland cereals.
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79
Yield Components and yield were recorded at harvest. Yield was obtained from 5
m2 harvest area and then adjusted to ton per hectare at 14% moisture content. Five
plants per plot were used to characterize the yield components.
Soil matric potential was measured by tensiometers' installed at 10 cm depth,
whereas perched water table was measured with piezometer inserted at the 75 cm
depth. Daily rainfall data were taken from a weather station located 50 m away from
the experimental site.
Experiment 2. Study on the effect of soil-water stress at reproductive stage on
grain yield in walik jerami rice.
Field experiments were conducted at the Jakenan Experimental Station during
1991 and 1992 dry seasons. A split plot design with three replications was used for
both experiments. In 1991, five rice genotypes IR 393557-71-1-1-2-2, IR39422-181-22,
lR459129-1-22,lR48563-11-2-2-3 and Dodokan were used as subplot and three levels of
soil water stress as the main-plot factor. The soil moisture stress treatments were : a)
continuous field saturation, b) no irrigation from panicle initiation(PI) to heading, and c)
no irrigation from heading to maturity. In 1992 rice, the genotypes used were:IR36,
IR39357-71-1-1-2-2, S969b-265-1-4-1 and S400b-55-2.
The 21-d-old seedings were transplanted at 2 seedlings/hill with 20 x20 cm
spacing. Land was ploughed twice by hoeing and harrowing. The plot size was 4 x
5 m and each plot received Urea), TSP and KC1 with the rates 135 kg N, 45 kg P2O5
and 50 kg K2 O ha-1 , respectively. Nitrogen fertilizer (Urea) was applied in three
equal splits of 45 KgN ha-1 each, at transplanting/ at 3 weeks after transplanting,
and at panicle initiation. All of KC1 and TSP were applied as basal. Weeds, pests
and diseases were controlled intensively.
Observations were made for yield components and yield. Four plants per plot
were used to characterize yield components. Grain yield was obtained from 8 m2
harvest area and calculated as ton per hectare at 14% seed moisture content.
Soil moisture status at 0-15 cm depth was determined by gravimetric method
and expressed as per cent oven dry weight (%ODW).
Experiment 3. Greenhouse study on the effect of soil water stress at different
growth stages on the yield of lowland rice.
This experiment was conducted at the Sukamandi Experimental Station during
1989 dry season using Ultisol soil with silty clay texture. A split plot design with four
replications was used. Four levels of soil water stress were used as sub-plot factor i.e.:
a) continuous flooding, b) water stress imposed from 10 DAT to panicle initiation, c)
water stress imposed from panicle initiation to heading and d) water stress imposed
from heading to maturity. Two varieties were used as main plot factor i.e. a) IR64 and
b) Ciliwung. The soil moisture stress was allowed to reach -50 cb and measured by
tensiometer at 10 cm depth. The water stress was imposed by not watering the pots.
After reaching the stress level, pots were reirrigated until field saturation. The 21-d old
seedlings were transplanted to each pot with 4 hills per pot.
82
Grain yield data were obtained from each pot and expressed in gram per hill
at 14% moisture content.
Crop data were analyzed using the methods for the split plot design and Duncans
Multiple Range Test was employed if F-test showed significance.
RESULTS AND DISCUSSION
The results of the effect of time of flooding on the yield components and grain
yield for gogorancah rice are presented in Table 1. Panicle length, 1,000-grain weight
and grain yield were affected by time of flooding. Flooding at four to six weeks after
seeding was important period to achieve higher yields. There was a tendency of
decline in grain yield when the flooding was applied more than six weeks after
seeding. Spikelet number per panicle and filled-spikelet percentage were not affected
by time of flooding treatments. There were significant differences in spikelet number
per panicle, filled- spikelet percentage and 1,000- grain weight but not in grain yield
between Dodokan and Cikapundung cultivars.
The decline in yield at, or above seven weeks after seeding might be due to
low supply of water and nutrients particularly nitrogen to the crop. Ismunadji (1985)
stated that the form and availability of nutrients are directly related to moisture
supply in the soil.
The effect of time of flooding on the mot mass density for the gogorancah rice
is illustrated in Figures 1 and 2.
Fig. 1 :
Root growth of Dodokan variety at the 0-5 cm soil depth as affected by time of flooding,
Jakenan, 1988-89 WS.
83
Fig. 2 :
Root growth of Cikapundung variety at the 0-5 cm soil depth as affected by time of flooding,
Jakenan, 1988-1989 WS.
Table 1.
Treatments
Cultivars (C)
Dodokan
Ci kapund ung
Time of flooding (F)
Weeks after seeding
(WAS)
4 WAS
5 WAS
6 WAS
7 WAS
8 WAS
9 WAS
CV (C):
CV (F):
*
Yield components
Spikelet/
panicle (nos)
Filled
spikelet
(%)
1,000
grain wt
(g)
Panicle
length
(cm)
84.3 b
119.4 a
75.0 a
69.5 b
24.7 a
23.5 b
22.3 b
23.1 a
105.4 a
98.3 a
103.0 a
105.1 a
101.5 a
97.6 a
74.4 a
73.5 a
70.7 a
71.9 a
71.1 a
72.0 a
5.3%
7.8%
4.7%
4.5%,
25.1 a
24.5 ab
24.2 abc
23.8 bc
23.3 c
23.6 a
3.3%
3.8%
23.4.a
22.9 ab
22.9 ab
22.4.b
22.3 b
22.3 b
3.9%
3.3%
Grain
yield
(t/ha)
4.17 a
3.95 a
4.54
4.24
4.10
4.05
3.81
3.63
a
ab
ab
bc
bc
c
18.9%
10.0%
There was gradual increase in root mass density from 37 to 88 days after
seeding. The rooting characteristics were influenced by water regime and type of
cultivars. Longer dry period relative to wet period tended to increase the root mass
density at 0-5 cm depth.
Apparently Cikapundung had higher root mass density at 0-5 cm depth than
Dodokan, particularly when flooding was done at nine weeks after seeding. Rooting
depth of both the varieties was deepest when grown under dryland condition.
An aerobic soil favours deep root growth and rice varieties differ markedly in
their rooting habits, both laterally and vertically (Yoshida and Hasegawa, 1982). The
fluctuations in soil matric potential and piezometric water level during gogorancah
growing season are illustrated in Figures 3 and 4.
Fig. 3 :
Fluctuation of the soil matric potential at the 10 cm soil depth and piezometric water level
for the two time of flooding of Dodokan. Jakenan, 1988-1989 WS.
High fluctuation in soil matric potential occurred from 25 to 40 days after seeding
and was subsequently relatively stable. The piezometic water level was close to the soil
surface from 45 days after seeding till harvest.
The dynamics of soil matric potential and the depth of piezometric water level
are partly controlled by rainfall. Weekly rainfall well distributed was from 45 days
afterseeding until harvest (Fig.5) and therefore, the soil matric potential and perched
water table were found stable.
85
Fig. 4 :
Fluctuation of the soil matric potential at the 10 cm soil depth and piezometric water level
for the two time of flooding of Chillwung, Jakenan, 1988-1989 WS.
The effect of rice genotypes and soil water stress at the reproductive stage on
the yield components and yield are presented in Tables 2 to 5. The results of genotype
x water stress interaction on the number of spikelets per panicle are presented in
Table 2. Dodokan and IR3922-18-1-2-2 were found to have stability in number of
spikelets per panicle under reproductive moisture stress.
Table 2.
Treatment
Rice genotypes (C)
Primordia
to heading
Heading to
maturity
Spikelet/panicle (no)*
IR393537-71-1-1-2-2
IR39422-18-1-2-2
IR45912-9-1-2-2
IR48563-11-2-2-3
Dodokan
CV (C):
CV (S):
86
96.7 a
69.4 b
51.7 c
56.3 c
80.5 b
74.4 b
99.9 a
60.7 c
68.4 bc
89.9 a
78.1 a
82.7 a
62.0 b
64.0 b
75.9 a
12.07%
9.3%
In a column, means followed by a common letter are not significantly different at the 5% level by
DMRT
The number of spikelets per panicle were shown to be reduced by reproductivephase drought (De Datta, 1981). The effects of moisture stress at the reproductive stage on
the yield components and yield are presented in Table 3.
Table 3.
Yield components and grain yield as affected by rice genotypes and soil
water stress at reproductive stage, Jakenan, 1991 DS.
Treatment
Yield Components*
Filled
spikelets (%)
1,000 grain
wt (g)
Grain yield
(t/ha)
65.8 a
1.96 a
0.97 b
0.36 c
PI to heading
Heading to maturity
Rice genotypes (C)
47.8 b
24.7 c
26.6 a
26.0 a
23.1 b
IR393537-71-1-1-2-2
51.3 a
25.9 a
1.20 a
1.19 a
IR39422-18-1-2-2
52.0 a
22.8 b
IR45912-9-1-2-2
40.8 b
25.6 a
0.99 b
IR48563-11-2-2-3
44.4 b
25.6 a
0.97 b
Dodokan
42.1 b
26.3 a
1.13 ab
CV (S) :
CV (C) :
16.0%
13.4%
5.3%
5.0%
In a column, means followed by a common letter are not significantly different at the 5% level by
DMRT.
During the growing season of walik jerami rice, occurrence of moisture stress
from panicle initiation to heading and from heading to maturity, significantly influenced
the filled spikelet percentage and grain yield. Moisture stress from panicle initiation to
heading and from heading to maturity reduced yield by 50% and 72%, respectively.
However, moisture stress from heading to maturity significantly influenced the 1,000
grain weight.
Moisture stress in the early reproductive stage (panicle initiation to heading)
resulted in a decreased grain yield through Education in number of spikelets.
Moisture stress during reproductive phase has been shown to reduce filled spikelet
number per panicle (IRRI, 1990). However, it had no influence on the panicle number
per hill, 1,000 grain weight and yield of walik jerami rice during the dry season of
1992 (data not shown).
87
Fig. 5 :
The genotype IR36 exhibited greater spikelet sterility under both the
continuous saturation and reproductive moisture stress situation. Apparently, soil
water stress from heading to maturity induced the spikelet sterility (Table 4).
Table 4.
Panicle initiation
to heading
Heading to
maturity
IR36
15.8 a
IR393537-71-1-1-22
13.3 a
23.9 ab
28.8 b
S969b-265-1-4-1
10.2 a
16.2 c
17.2 b
S400b-55-2
18.7 a
23.0 ab
19.5 b
CV (C) :
29.6%
CV (S) :
31.0%
88
The rice genotypes grown in 1992 walik jerami did not show any significant
differences in grain yield. Water stress treatments at reproductive stage had no effect
either. The average yield ranged from 2.2-2.6 t/ha. The genotypes IR36, however,
exhibited a tendency to yield higher (though not significant) than other breeding
lines (data not presented). The rainfall distribution was optimum during the
stress-treatment periods and hence the soil moisture status reached above field
capacity. This is considered as critical level of soil moisture for rainfed lowland rice
(Fagi, 1986), below which significant yield differences could be expected.
A comparison between two genotypes (grown in 1989) to moisture stress
tolerance revealed that the level of yield of Ciliwung was lower than IR64.
However, it was found to be more tolerant to moisture stress at various growth
stages. IR64 was more sensitive to moisture stress from panicle initiation to
heading than other phases of moisture stress, as evidenced by a drastic reduction
in grain yield (Table 5).
Table 5.
Cultivar (C)
IR64
Ciliwung
grain yield* (g/hill)
Continuous
flooding
20.82 a
17.26 a
18.82 a
15.71 a
PI to Heading
10.31 b
15.04 a
Heading to maturity
19.90 a
17.25 a
CV (C) :
3.0%
CV (S) :
10.7%
In a column, means followed by a common letter are not significantly different at the 5% level by
DMRT
CONCLUSION
In Pati district, the optimum time of flooding in gogorancah rice is in the
range of four to six weeks after seeding to obtain higher yield. The moisture stress
from panicle initiation to heading was found to be critical in walik jerami rice. The
genotypes Ciliwung and TR39357-71-1-1-2-2 were found, having potential to be
planted as Walik jerami rice.
89
REFERENCES
De Datta S K (1981) Principles and practices of rice production. John Wiley & Sons.
Fagi A M (1986) Efficient use of water for rainfed lowland rice. In : Progress in rainfed lowland rice. IRRI,
Los Banos, Philippines.
IRRI (1990) Program report for 1989. International Rice Research Institute., Los Banos, Philippines.
Ismunadji M (1985) Effect of sulphate application on the performanceof IR36 rice variety under submerged
and dryland condition. Indonesian Journal of Crop Science l(1) : 1-64.
Yoshida S, S Hasegawa (1982) The rice root system, its development and function. In : Drought resistance
in crops with emphasis on rice. IRRI, Los Banos, Philippines.
90
SUMMARY
Rice seedling establishment in wet direct seeding is erratic because farmers
sow seeds on the soil surface. Anaerobic- tolerant cultivars that grow vigorously
in flooded or water- saturated soil were identified. These cultivars developed
longer coleoptiles than non-tolerant controls, suggesting that they can transport
02 in the air or water to the seeds in anaerobiosis efficiently. The superiority of
tolerant cultivars in seedling survival and growth persisted even when grown in
the dark, although their growth increased under light. Temperatures between 15
and 35C had little effect on difference in seedling survival between tolerant and
control cultivars, although seedling growth of both types of cultivars was reduced
at 15C.
The growth of a tolerant cultivar in flooded soil was equivalent to that of a
control grown in drained soil. Thus, the use of anaerobic cultivars may make sowing
in flooded or water-saturated soil similar to that in freely drained soil. The potential
of its applicattion to rainfed lowland ecosystems was tested in Tarlac, Philippines,
during 1993 wet season. Rice seeds soaked in water for 24 h were dibble-sown into
water-saturated soil under zero tillage. There was little difference between
anaerobic-tolerant and control cultivars in crop establishment, presumably due to
soil dryness (there was no rainfall after seeding). The tolerant cultivars outyielded
the controls: the six anaerobic cultivars had a mean grain yield of 3.2 t/ha at 0-180
kg N/ha, whereas the control cultivars had 2.4 t/ha. Further agronomic and
physiological
studies
are
needed
to
adapt
anaerobic-tolerant cultivars
in
heterogeneous rainfed lowland soils.
INTRODUCTION
Crop establishment
A rice crop is established either by transplanting or by dry or wet direct
seeding. Transplanting is mostly practiced in irrigated lowland rice and is an
established technology. Dry seeding is also well studied and is practiced where
1 International Rice Research Institute, P.O. Box 933, 1099 Manila, Philippines
soil is dry. Rice plants adapted to irrigated lowland and upland ecosystems are bred
for transplanting and dry seeding, respectively, in Asia.
Wet seeding technology, however, has not been well developed. Germinated
seeds are sown on the surface of drained, water- saturated puddled soil or flooded
soil surface (water seeding). Crop establishment by wet seeding seems to be more
unstable and inconsistent than that by dry seeding. In dry seeding, seeds are covered
with aerobic soil and protected from birds, rodents, and rain splashing; in wet
seeding, seeds are sown on puddled soil surface and exposed to these disturbances.
The condition of the puddled soil surface changes according to intensity of puddling,
climate, and time after land preparation. In water seeding, seedlings not only drift
due to poor anchorage but are also destroyed by snails and fish.
Rice seedlings grown by wet seeding are exposed to soil anaerobiosis, although
this is not a constraint in transplanted and dry seeded rice. Because there are no
cultivars raised and adapted for wet seeding, farmers currently use cultivars
developed specifically for transplanting.
Anaerobic seeding is proposed to improve the wet seeding process (Yamauchi
et al., 1993a). Germinated seeds are sown under the surface of puddled soil with or
without standing water so that seeds are protected from the disturbances occurring
on the soil surface. In addition, seeds get better anchorage in water seeding. Because
seeds are exposed to soil anoxia, cultivars tolerant of anoxia are preferable for this
seeding method.
The process of crop establishment in wet seeding could be divided into survival
(which can be measured by percentage seedling establishment) and growth (which can
be characterized by leaf development, seedling height, and shoot and root weights).
Mesocotyl elongation is an important factor in raising the apical meristem to the upper
soil layer when seeds are sown deep into the soil (Takahashi, 1984).
Germplasm selection
A screening system to find rice germplasm suitable for anaerobic seeding was
developed (Yamauchi et al., 1993a). Seeds that germinated for 2 d were planted 25
mm deep in seedling trays. The trays were then submerged in water to a depth of
30-50 mm. Seedling establishment was evaluated by analyzing leaf development,
seedling height, and percentage establishment (survival) 15 d after planting.
Rice germplasm tolerant to soil anoxia were identified in traditional cultivars and
also in breeding lines with improved plant type. Screening for anaerobic-tolerant rice
germplasm was conducted using 256 accessions from conserved germplasm from the
International Rice Germplasm Center (IRGC) and 404 accessions from the International
Network for Genetic Evaluation of Rice (INGER), IRRI (Yamauchi et al., 1993 a). IRGC
germplasm represented broad genetic diversity, and INGER germplasm involved
desirable agronomic characters. Eight percent of the IRGC germplasm and 2% of those
from INGER were identified statistically as tolerant of soil anoxia. Among the anaerobic
germplasm were those from northeast India and Bangladesh which were adapted to
deepwater and early summer rainfed lowland cultures.
92
The yield potential of anaerobic tolerant cultivars with improved plant type
was equivalent to or higher than that of high-yielding local check cultivars when
grown under direct seeding. In yield trials at Los Baos, Philippines, the
anaerobic-tolerant cultivar IR52341-60-1-2-1 had the same high yield as check IR72
(8.4 t/ha). At Muoz, IR52341-60-1-2-1 yielded more than IR 72 (9.8 vs 7.0 t/ha). In
the Mekong Delta, anaerobic-tolerant cultivar BR1870-67-1-3 yielded 5.8 t/ha,
whereas check MTL103 yielded 5.1 t/ha during the 1993 wet season (Chau and
Yamauchi, 1994). IR52341-60-1-2-1 yielded 5.2 t/ha in Hanoi with check CN2 yielding
4.5 t/ha during the 1993 wet season (Chuong and Yamauchi, 1994).
Wet seeding in the rainfed lowland ecosystem
Because soil water conditions in rainfed lowland fields vary according to
location in toposequence, rainfall, and time after the onset of the wet season, farmers
choose dry seeding, wet seeding, or transplanting in order to establish the crop
successfully and economically. Drought is a serious constraint in rainfed lowland
rice culture. Early planting by direct seeding advances the plant developmental stage,
thus avoiding drought at the termination of rainfall at the end of the wet season.
Direct seeded plants have both seminal and nodal roots, whereas transplanted plants
have only nodal roots. In addition, seminal roots might develop deeper than nodal
mots at the early growth stage. Thus, direct seeded rice might be more tolerant of
drought at the early stage than transplanted rice.
Compared with dry seeding, wet seeding is less commonly practiced in the
rainfed lowland fields, presumably due to the instability of crop establishment.
Success in wet seeding depends on the characteristics of the dry-to-wet transition
period at the onset of the monsoon and on adequate surface drainage (Morris and
Zandstra, 1979). Water seeding is practiced where drainage is difficult. Introducing
anaerobic seeding in the rainfed lowlands would stabilize crop establishment in wet
seeding, and thus may interest farmers to follow direct seeding.
Seedling growth of anaerobic-tolerant cultivars in flooded soil is characterized
here in terms of light and temperature, and in comparison with growth in drained
soil in controlled environments. We also studied the crop establishment and grain
yield of rice plants in rainfed lowland fields with water- saturated soil under zero
tillage to identify the contraints in adoption of anaerobic seeding.
MATERIALS AND METHODS
Anaerobic seedling growth under continuous light and darkness
Seeds of three anaerobic-tolerant cultivars (CO25 [percentage germination,
93.4%; rate of germination, 1.00], ASD1 [99.6; 1.001, and Taothabi [98.3; 1.001 and
three control cultivars IR36 [96.5; 1.00 IR50 [95.2; 0.951, and IR74 [91.7; 0.99]) were
germinated in petri dishes for 2 d at 30C, and then sown in sieved (mesh 40) upland
(Maahas clay) soil placed in a container (300 x 240 x 110 mm). Rate of germination
was measured according to Krishnasamy and Seshu (1989). The container was filled
with soil up to a height of 80 mm. Seventeen seeds were sown for each cultivar per
94
container. The soil was then submerged under 30 mm of water. Eight containers
were placed in a growth chamber (30C, 70% relative humidity) under continuous
darkness or light (30 k lux). Plants in four containers (replications) were sampled 7
and 11 d after seeding.
Seedling growth was characterized by measuring emergence score (0=no
emergence, 1=coleoptile emerged, 2=lst leaf emerged, 3= 2nd leaf emerged, ...) ;
seedling height; and mesocotyl length of the individual plant; and shoot, mot, and
seed weights as a bulk sample per replication after drying at 80C. Percentage of
seedlings established was calculated to estimate seedling survival.
Effect of temperature on anaerobic seedling growth
Seeds of six cultivars (CO25 [percentage germination, 93.7%; rate of
germination, 0.991, ASDl [l00; 1.001, Taothabi [97; 0.941, IR 36 [95.5; 0.981, IR50 (93.3;
0.931, and IR74 [91.3; 0.991) were germinated and planted using the method described
above. The containers were placed in temperature-controlled glass rooms (outdooor
growth chamber) (under natural light) using the set of 15, 20 and 30C or 25, 30, and
35C. One container represented one replication. Four containers (replications) were
placed in each glass room. Plants were sampled 14 d after seeding and their growth
and survival analyzed.
Seedling growth in flooded and drained soils
Seeds of IR36 (percentage germination, 96.8%; rate of germination, 0.98) and
Taothabi (98.9; 0.97) were similarly germinated and sown, except that the plastic
containers were replaced by 3.8-liter pots. The soil in the pot was flooded at a depth
of 5 cm or freely drained, Seventeen seeds of the cultivar were sown per pot. The
pots were placed in a temperature- controlled (29/2lC day/night) glass room
(phytotron) under natural light . Plants were sampled 4, 6, 8, 10, 12, and 14 d after
seeding and their growth was measured. Root length (from the tip of seminal root
to the base of plant) was measured. The experimental design was a randomized
complete block with four replications (pots).
Performance in rainfed lowlands
Seeds of six anaerobic-tolerant cultivars (IR341996-50-2-1-3 [percentage
germination, 99.3; rate of germination, 1.00], IR52341-60-1-2-1 [100; 1.00],
IR50363-61-1-2-2 [99.3; 0.991, IR 31802-48-2-2-2 [100; 1.00], BR1870-67-1-3 [98.5; 0.89],
and BR736- 20-3-1 [100; 1.00]) and two controls (PSBRC10 and IR72) were
dibble-sown under zero tillage in a rainfed lowland field at Masalasa, Tarlac,
Philippines, during 1993 wet season. All seeds were produced during 1993 dry
season; dormancy was broken by keeping seeds at 50C for 5 d.
After dry-season harvest, the remaining straw was removed from the field.
The seeds were soaked in water for 1 d, then dibbled manually at 5-10 seeds per
hole at a depth of 25-30 mm at 20x 20cm spacing. Soil was saturated with water at
sowing and dried after 1 wk. Weeds were removed by hand. Fertilizer was applied
at the rate of 0, 60, 120, and 180 kg N/ha in four splits (15 and 35 d after emergence,
95
on seedling
affected by
weight and
at 30C.
Fig. 1.
Rice seedling growth in flooded soil under continuous light and darkness at 30C. Seedling
growth was measured at 7 and 11d after seeding. Bars having a common letter (under light
or dark condition at 7 or 11d) are not significantly different at the 5% level by DMRT.
Number denotes cultivar names: 1=IR36; 2=IR50; 3=IR74; 4=CO25; 5=ASD1; 6=Taothabi.
The shaded and closed bars indicate the light and dark conditions, respectively.
97
by seed rate and seedling survival) and single-seedling weight. Therefore, crop
establishment may be low at lower temperatures and the highest at 30C.
Fig. 2.
98
Effect of temperature on rice seedling growth under natural light condition. Growth was
measured 14d after seeding. Experiments were conducted 2 times with the temperature
combinations of 15, 20, and 30C (closed symbols) and 25, 30 and 35C (open symbols).
Values are the means of control (IR36, IR50. and IR74) ( ) and anaerobic tolerant cultivars
(CO25, ASD1, and Taothabi) (O). The difference in each character between control and
tolerant at each temperature was significantly different at the 1% level (even at 15C for
seedling height and shoot dry weight).
Fig. 3.
Seedling growth and seed weight of rice cultivar IR36 (control) ( ) and Taothabi (anaerobic)
(O) in flooded (closed) and drained soils (open). Values having a common letter at each
day are not significantly different at the 5% level by DMRT.
99
unidentified stresses. The results suggest that stress tolerance in rainfed lowland rice
is important in increasing the yield.
Table 1.
Cultivars
Crop establishment**
Height
(mm)
Plant
density
Shoot
dry wt.
Mesocotyl
length
(mm)
0
(no./m 2 )
(g.m 2 )
IR4199650-21-3
124bcd
10.5
216
5.4
IR5234160-1-2-1
147ab
12.9
169
IR5036360-1-2-2
116cd
8.1
IR31802
48-2-2-2
143abc
BR187067-1-3
N levels (kg/ha)
60
120
180
3.0a
3.3ab
3.7a
3.3bc
5.4
2.4abc
2.9abc
3.3ab
4.0ab
170
5.0
2.1bcd
2.9abc
3.5a
3.4bc
12.3
181
7.3
2.0cd
3.5a
3.6a
3.8ab
96d
7.2
149
5.1
2.8ab
3.0abc
3.7a
3.3bc
BR73620-3-1
102d
9.3
191
7.1
2.0bcd
2.9abc
3.5a
4.6a
PSBRC10
(Control)
169a
12.6
177
6.9
1.1e
2.6bc
3.1ab
2.9c
IR72
(Control)
123bcd
12.5
176
6.6
1.5de
2.3c
2.7b
2.9c
In a column, means followed by a common letter are not significantly different at the 5%
level by DMRT.
**
RESEARCH NEEDS
The results presented here demonstrated that anaerobic-tolerant cultivars had
superiority in seedling establishment over a wide range of environmental conditions
: light, temperature, drained/flooded soil. In addition, they were more tolerant to
the unidentified stresses in the rainfed lowland field in the Philippines, producing
more grains than the non-tolerant controls. Introducing anaerobic cultivars into
rainfed lowlands would increase not only the stability of wet seeding but also the
grain yield. Systematic research is needed to develop wet seeding technology in the
rainfed lowlands, in as much as crop establishment and grain yield are influenced
101
102
SUMMARY
Rice accumulates significant amount of carbohydrate in different plant parts
especially in the culm which has an important role when plant experiences drought
and flooding at various stages of crop growth. Carbohydrate status of rainfed
lowland rice vis-a-vis mechanism of submergence tolerance at seedling stage,
drought at flowering stage and shade at reproductive stage has been discussed with
experimental evidences. Tolerance to submergence at seedling stage, and drought
at flowering was closely associated with higher levels of carbohydrates (total water
soluble carbohydrate and starch). Hydrolytic enzyme, alpha amylase activity had
positive correlation with seedling survival at initial stage of submergence and was
negatively correlated during submergence. Yield of rice under water deficit at
booting or flowering was directly related to carbohydrate content, apparent
translocation rate (ATR) and apparent contribution rate (ACR). Chemical desiccation
method for screening flowering stage drought resistance has been discussed.
Histochemical analysis technique for screening for flowering stage drought tolerance
in rice has also been discussed. Adaptive features of wild rice species Oryza
australiensis in relation to drought has been discussed which include reduced tiller
and leaf area, more carbohydrate in culm and rhizomes.
INTRODUCTION
Rainfed lowland rice represents one of the major rice ecosystems of South
and South-East Asia. About one-fourth of the world rice area falls under this category
and contributes 19% of total rice grain yield. In India, approximately 14 million
hectare rice lands are rainfed lowland, of which major area (60%) comes from Eastern
India. Eastern Uttar Pradesh constitutes about 66% of rainfed lowlands in the state
(Table 1).
The productivity of rainfed lowland rice is low and averages around 1.0-1.5
t/ha which, though is higher than upland and deep water rice, but much lower than
that of irrigated ecosystem. Productivity of rainfed lowland is about 48% of value
product per hectare of irrigated rice (ADB, 1989). Among the general causes of poor
productivity of rainfed lowland are (i) poor establishment of seedlings due to
1
partial to complete submergence (Chaturvedi et al., 1993; Vergara, 1985), (ii) late
season drought/anthesis drought (Chaturvedi and Ingram, 1989, (iii) low light stress
in wet (Kharif) season (Murty and Sahu, 1987; Venkateswarlu and Srinivasan, 1978),
(iv) accumulation of toxic substances in ill-drained soils (Ponamperuma, 1984), (v)
soil factors such as salinity/alkalinity and zinc deficiency (Rao and Biswas, 1979),
(vi) cold stress at panicle initiation (Nanda and Coffman, 1979) and (vii) severe
occurrence of diseases and pests etc. Among the causes mentioned above,
submergence and drought are the most prevalent stresses to reduce yield, and
account for about 70% and 25% of yield loss, respectively in rainfed lowland rice
area (Widawsky and O'Toole, 1990). Thus evaluating and screening cultivars for
submergence at initial stage and for drought at later stages of crop growth become
an important pre-requisite to improve overall productivity of rainfed lowland rice.
Besides, there is a need to screen for salt, shade and cold tolerance for
location-specificity.
Table 1 .
Region
Rainfed Lowland
(000 HA)
(000 HA)
Rice Area
World
146400
37950
Asia
131000
36200
India
41000
14601
Eastern India
24000
8880
Uttar Pradesh
5500
2245
Eastern U.P.
2661
1415
A REVIEW
Carbohydrate status and stress tolerance
Most of the carbohydrates are produced by green leaves and other green
tissues of plants and are either translocated as sugar or stored temporarily as sugar,
starch or fructans. Later in the dark, sugar is resynthesized and translocated from
the leaf. In most plants, sucrose is translocated via phloem to carbohydrate sinks for
cell growth, metabolism, respiration or storage. Growing cells act as carbohydrate
sinks but later they may act as source when they become photosynthetically active.
At subcellular level chloroplast may act as carbohydrate source and mitochondria
of same cell act as sink (Kozlowski, 1992).
Growth and grain yield depend on both current photosynthesis and stored
assimilates. If current photosynthesis is limited by environmental stresses such as
water deficit (Marshall et al., 1989 Bunce, 1982: Blum et al., 1983; Chaturvedi and
Ingram, 1988), low temperature (Hilliard and West, 1970; Chatterton et al., 1972),
nutrient deficiency (Lenhert et al., 1979), or submergence (Setter et al., 1989 a and b;
Chaturvedi et al., 1993), the remobilization of stored assimilates is accelerated and
plant is forced to depend on stored assimilates. Stored assimilates also enhance
recovery of plants after stress (Wardlaw and Eckhardt, 1987).
Stored carbohydrates have important role in metabolism, growth,
development or stress tolerance, defense and survival (Kozlowski, 1992). It is
therefore, also called reserve carbohydrate (RC) and is particularly sensitive to late
season stress and management practices. It is also essential for survival when
photosynthesis is low or is stopped. Among stored carbohydrates, starch is the most
important reserve carbohydrate. Starch accumulates whenever high level of sugars
build up and starch is transformed into sugars when sugars are low (Kozlowski and
Keller, 1966). Among soluble carbohydrates, sucrose is the major translocable and
storage carbohydrate. Reserve carbohydrates are also important in preventing
invasion by certain plant pathogens and insect attacks.
Carbohydrate status and submergence tolerance
Rice crop suffers from partial to complete submergence in rainfed lowland
areas. Complete submergence occurs during short- term flash flooding, varying upto
15 days, near river valleys and riverlets. Characteristics of flood water vary at
different locations and the nature depends upon light intensity under water, turbidity
and concentration of dissolved gases such as O2, CO2 and C2H4. Partial submergence
of rice is quite common in low lying areas.
Complete submergence of rice leads to low concentration of carbohydrates
and reduced growth and finally death of tissues (Palada and Vergara, 1972). It is
partly due to low rate of photosynthesis (Trought and Drew, 1980; Rai and Murty,
1979), which is directly associated with slow diffusion of CO2 in water (Setter et al.,
1989 a). Other factors which contribute to slow growth are (i) acceleration of
photorespiration, (ii) anaerobiosis leading to breakdown of CHO and carbon loss as
CO2 and ethanol, (iii) high ethylene production disturbing membrane integrity and
105
chlorosis, (iv) leakage of CHO, amino acids and ketones, (v) ABA accumulation and
decreased supply of cytokinins which lead to stomatal closure and (vi) reduction of
RUBP carboxylase activity. Complete submergence reduces level of carbohydrates
but in partial submergence the reduction depends on proportion of leaves below
the water (Setter et al., 1989). Submergence reduces concentration of soluble sugars
and starch in all plant parts of rice by 4-12 fold (Setter et al., 1989).
Amount of carbohydrate in plant parts correlates positively with submergence
tolerance (Palada and Vergara, 1972; Emes et al., 1987; Chaturvedi et al., 1993). Older
seedlings which have greater submergence tolerance, also contain more carbohydrate
(Vergara, 1985; Chaturvedi et al., 1993). Decreasing seedbed soil nitrogen level
increases seedling carbohydrates and submergence tolerance as well (Vergara, 1985).
Emes et al. (1987) observed that under submerged condition, the culm of
submergence-tolerant varieties accumulated higher levels of starch and at a more
rapid rate than did varieties with low submergence tolerance. Culms of submergence
tolerant plants contained starch even after being submerged for 7 days, whereas the
starch reserve of submergence susceptible varieties was exhausted during the same
period. Plant may use reserve starch stored in the culm during submergence, and
the ability to survive for longer periods may be related to continued availability of
carbohydrates. Chaturvedi et al. (1993) showed that levels of total water soluble
carbohydrates and starch were positively correlated with tolerance to
complete/partial submergence. Tolerant genotypes had lower rate of reduction of
CHO during submergence and higher recovery after 30 days. Slow rate of reduction
in tolerant genotypes makes the substrate available for regeneration of growth after
releasing the submergence. Alpha- amylase activity was also higher before
submergence and after recovery in tolerant genotypes. In contrast, under partial
submergence condition, leaf sheath plays the major role in carbohydrate distribution.
High carbohydrate depletion rate (CDR) in leaf sheath was observed in tolerant than
susceptible genotypes under partial submergence condition.
Submergence for 3 days under continuous light led to mobilization of 65% of
the starch from the regions of rice internodes which has been formed prior to
submergence and disappearance of starch was accompanied by a 70-fold
enhancement of amylolytic activity (Raskin and Kende, 1984). Submergence also
caused 26-fold increase in the translocation of newly synthesized photosynthate
assimilate from leaves to internodes and younger regions of culms. Tops of rice
contain significantly larger amounts of starch than those of submerged plant
(Yamaguchi, 1973). Amylase activity increases in rice leaves and internodes during
submergence (Yamaguchi and Sato, 1963).
Carbohydrates status and drought tolerance at reproductive stage
Substantial quantities of carbohydrates are accumulated in different parts in
rice (Perez et al., 1971) which are of paramount importance when plant experiences
water deficit at flowering (Rahman and Yoshida, 1985; Chaturvedi and Ingram, 1988,
1989). Pre-anthesis assimilates stored in stem and leaf sheath tissue appear to
contribute substantially to grain filling under water stress (Akita, 1987; Cock and
106
non reducing sugars and starch) are markedly reduced in all plant parts. Total soluble
N and amino N are enhanced in susceptible varieties. Higher sterility is ascribed to
disturbed N metabolism and accumulation of soluble N in the panicle which is toxic
to normal grain setting (Murty et al., 1983).
EXPERIMENTALRESULTS
Submergence tolerance
Submergence tolerance in rice is not governed by a single factor. Our previous
results (Chaturvedi et al., 1993) indicated that elongation and submergence tolerance
are two independent traits. Attempt has been made to combine these two traits in
one genotype (Raysaha et al., 1993). Submergence tolerance is a dominant character
and additive gene effects appear to be operative in the control of this trait (Mohanty
and Khush, 1985).
From our three years (1991-1993) data, it is clear that initial high carbohydrate
level in the tissues is one of the adaptive traits of submergence tolerance. Higher
levels of initial carbohydrate (water soluble sugar and insoluble starch) act as buffer
stock and its continued availability during submergence is critical for the survival
and growth of rice. Metabolic energy needed by the plant during submergence is
primarily supplied from stored carbohydrates present in the tissue.
Tolerant genotypes had higher total initial carbohydrate content and starch
before submergene and had lower rate of depletion during submergence than the
susceptible (Fig. 1 & 2). Susceptible genotypes had poor accumulation of CHO and
showed faster rate of depletion during submergence. Slow rate of depletion of CHO
in tolerant genotypes seems to be one of the adaptive traits to withstand submergene
due to continuous availability of substrate for energy for a longer period. In
susceptible plants, drastic reduction of CHO leads to high rate of anaerobic
fermentation and production of ethanol at toxic level. Setter et al. (1989b) also
proposed the accumulation of end products to toxic levels in their model on response
of rice to submergence.
Carbohydrate status of plant changes with the type and nature of
submergence, plant age and exposure to water phase (Setter et al., 1989b). In one of
the experiments, plants were completely submerged and both shoots and roots were
under water, carbohydrate depletion occurred and it was more in susceptible than
in the tolerant genotypes.
Tolerant genotypes had higher recovery of total carbohydrate and starch 30
days after releasing submergence stress than the susceptible ones. The alpha amylase
activity was higher before as well as after releasing stress (Fig. 3). Higher alpha
amylase activity in leaf and culm seemed important for longer submergence duration
tolerance in tolerant genotypes. In general, older seedlings (45- day old) had better
survival than younger seedling (15, 30-day old) and it was due to higher level of
stored CHO in older seedlings. Vergara (1985) also observed higher level of CHO
before submergence which decreased at low rate during submergence.
108
Fig. 1.
109
Fig. 2.
110
Changes in total water soluble sugar (WSS) in tolerant and susceptible genotypes in complete
submergence (15 days). Pooled data of 1992-93.
Fig. 3.
111
Contrarily, in deep water rice, Emes et al., (1987) reported that culms of
submergence tolerant varieties maintained higher levels of starch, and its accumulation
was more rapid than the less tolerant type. Raskin and Kende (1984) working with deep
water rice variety-Habibganj Aman from Bangladesh, found that starch disappearence
was acompanied by 7-fold enhancement of amylolytic activity. Internodes of air grown
rice plants contained large amounts of starch (about 10% DW) which disappeared upon
submergence in water under continuous light. This indicates that carbohydrate status
and its remobilization alter with the status of plant in water.
Correlation studies showed that survival of rice was positively correlated with
the level of total water soluble sugar and starch in the culm, before submergence,
after submergence and after recovery (Table 2). Strong positive correlation was
found in shoot (culm) after submergence stage rather than before and after recovery.
There was a negative correlation between alpha amylase and survival in both mot
and shoot after submergence.
Table 2 .
Character
Correlation values between survival and starch, survival and total water
soluble sugar and survival and alpha amylase activity in different plant
parts. 30 d old seedling, 11 d submergence. Pooled data of 1992-1993.
Relationship
r value*
B.S.
A.S.
A.R.
N.S.
0.95**
N.S.
0.99**
0.99**
0.96**
N.S.
O.80**
N.S.
0.76**
N.S.
0.92**
0.85
0.94
-0.90
-0.99
N.S.
N.S.
Root
Shoot
Drought tolerance
Rice accumulates significant amount of carbohydrates before heading and
there exists wide variability among genotypes. Improved rices have higher contents
of total non-structural carbohydrate at heading than do the low yielding traditional
varieties (Weng et al., 1986). Previous experiments have shown that when water
deficit occurs during flowering or grain filling, grain growth derives a greater
112
Plant Parts
Tillering
Booting
Flowering
0.23
0.42*
0.78**
Leaf sheath
0.20
0.49*
0.51*
Culm
0.14
0.19
0.46*
0.21
0.47
0.85**
Leaf Sheath
0.20
0.41*
0.66**
Culm
0.20
0.46*
0.52**
and **
Yield was inversely Correlated with apparent translocation rate (ATR). Among
cultivars, yield was directly correlated with maximum ATR under stress condition,
and cultivar IR-46 maintained better translocation under stress than did the other
cultivars (Table 4).
One of the goals of plant breeders is to produce germplasm with stable yields,
and a measure of yield stability is the slope of ATR vs yield regression, the smaller
the slope, the more stable the yield. Variety IR-64 had the greatest slope and high
crop susceptibility (cs) factor.
113
Table
4.
contribution
rate
Cultivars
Time of
water deficit
ATR
ACR
Yield
g/plot
Crop
sucesptibility
IR64
Control
Tillering
Booting
Flowering
0.10
0.17
0.24
0.29
0.13
0.21
0.34
0.39
19.4
16.7
12.8
11.9
0.13
0.34
0.38
IR46
Control
Tillering
Booting
Flowering
0.14
0.21
0.31
0.42
0.16
0.26
0.44
0.51
22.3
19.2
15.7
14.7
0.06
0.22
0.23
Mahsuri
Control
Tillering
Booting
Flowering
0.08
0.15
0.19
0.25
0.11
0.16
0.26
0.32
15.9
13.2
11.1
09.8
Salumpkit
Control
Tillering
Booting
Flowering
0.08
0.13
0.22
0.31
0.11
0.16
0.31
0.42
14.3
11.8
10.1
11.1
0.17
0.29
0.22
IR54
Control
Tillering
Rooting
Flowering
0.12
0.19
0.27
0.36
0.14
0.22
0.37
0.40
20.3
18.3
13.8
11.8
0.13
0.25
0.31
0.04
0.05
0.6
0.05
LSD 0.05
0.17
0.31
0.39
In the histochemical studies also, it was observed that tolerant genotypes had
higher level of starch localization at flowering in non-stress environment than
susceptible genotypes. Histochemical observation of CHO localization was very
close to the observation recorded by quantitative method (Tables 5-7). Under stress,
carbohydrate is remobilized relatively faster in tolerant than in susceptible genotypes
as also evident by carbohydrate depletion rate (CDR) (Table 6). It is evident from
Table 7, that yield of tolerant cultivar IR 46 was relatively higher under stress
condition than that of susceptible variety IR 52. Higher level of sugar acts as osmotic
agent to maintain higher leaf area and dry weight. Poor yield in susceptible cultivar
IR 52 was mainly due to its poor storage of CHO and poor translocatory behaviour
which was reflected in its poor dry weight and, leaf area, and more unfilled spikelets
(64%) in comparison to tolerant ones (30%).
114
Table 5.
Cultivars
After recovery
IR-46
3000
1390
580
N-22
2230
900
420
IR52
2300
1200
840
470
281
136
LSD 0.05
Table 6.
Cultivars
rice
genotypes
IR-46
40
80
N-22
38
64
IR52
28
34
LSD 0.05
14
Table 7.
Cultivars
Grain yield
(g/pot)
C
WD
IR46
20.6
14.0
105
N-22
16.4
12.4
IR52
17.9
9.0
LSD 0.05
1.1
Spikelet/
panicle
WD
Individual
grain weight (mg)
C
WD
Unfilled
grain (%)
C
WD
83
23
20
16.5
30.5
87
71
19
18
20.0
30.5
107
87
24
19
19.7
3.0
1.1
63.8
3.1
in leaf blade, while Oryza australiensis had higher accumulation of CHO in culm and
leaf sheath.
Table 8
Wild rices
Water treatment
Leaf
CHO (mg/pot)
Culm
Oryza
australiensis
Control
Stress
1500
1050
3600
2340
1080
646
40
60
Oryza officinalis
Control
Stress
1800
1530
3000
2400
780
585
30
35
Oryza nivara
Control
Stress
2400
2160
430
2560
2048
570
750
600
375
40
48
LSD 0.05
Root
CDR
mg/pot/day
All wild species showed rhizomatous characters, full of starch, but Oryza
austruliensis had higher starch localization and remobilization than other wild species
as evident by histochemical and quantitative analysis. This was reflected in higher
yield under stress condition. Oryza nivara, which produced higher dry matter, did
not produce enough yield under stress. Among 3 wild species, Oryza australiensis
showed more adaptive traits like reduced tiller and leaf area, and more carbohydrate
content before stress and faster remobilization during stress. Higher intensity of
starch localization in rhizomes is a strong indicator of tolerance behaviour under
stress.
In one of the experiments, a chemical dessicant was used as an agent for
simulating drought to identify genotypes for drought tolerance at flowering.
Carbohydrate depletion rate in vegetative tissue gives a relative estimate of CHO
remobilization to panicles. Except for roots, IR 46 had the highest CDR in all plant
parts. Thus, drought tolerant IR 46 had both the traits of accumulation and
remobilization of CHO. Both traits together confer tolerance to water deficit or
chemical dessication at flowering. Cultivar N22 had the highest CDR in its roots.
High root and culm CDR may explain renewed tiller production at the expense of
grain production. Our results are in close agreement with the work on wheat done
by Blum et al, (1983) and in rice by Chaturvedi and Ingram, (1991). Submergence
tolerant cultivar had minimum CDR value, and therefore showed that faster
remobilization is not a feature of submergence tolerance.
All cultivars responded similarly to water deficit and chemical dessication
with respect to CHO distribution, leaf area, dry matter, and yield and yield
components. Injury index by drought was more closely related to injury index by
chemical dessication at 5 days after panicle emergence (DAPE) than at 10 days before
116
panicle emergence (DBPE). The results showed that chemical dessicant had an effect
on rice yield similar to that of water deficit, with the common effect, that of forcing
the plant to rely on previously accumulated assimilates for grain growth. Chaturvedi
and Ingram (1991) also found that low concentration of atrazine (0.05 kg a i/ha) in
a non-stress environment, was a potential simulator of drought.
Shade tolerance
In general, shade reduced CHO in all plant parts relative to unshaded control.
The greater reduction in CHO however, was observed when shaded at 5-20 DAPE
than 10-25 days after panicle initiation (DAPI) in both wet and dry seasons.
Reduction in CHO by shading was also reported by Eaton and Ergle (1954), and
Kemp (1981).
Table 9 .
Stage
Treatment
Leaf blade
Culm
10 DAPI
WET
0.79**
0.72
0.53
DRY
0.75*
0.73
0.62
WET
0.45
0.34
0.59
DRY
0.35
0.36
0.62
WET
0.78**
0.76*
0.52
DRY
0.76*
0.80**
0.61
WET
0.48
0.29
0.09
DRY
0.22
0.25
0.02
WET
0.49
0.07
0.37
DRY
0.13
0.07
0.10
25 DAPI
5 DAPE
20 DAPE
HARVEST
Leaf sheath
Grain yield was positively correlated with CHO in leaf blades and culm at 10
DAPI and 5 DAPE (Tables 9 and 10). The culm was the greatest source of translocated
CHO for grain growth is supported by ACR. The results indicate that under water
deficit, grain growth depended on previously accumulated assimilates. Grain yield
and ATR were negatively correlated, but correlation was positive with maximum
ATR value. Our results are similar to those of Reyniers et al. (1982) who also reported
higher ATR values under water deficit. Translocation in supplying CHO to grain
increases when plant experiences stress (Austin et al., 1980). Upto 31% of rice grain
yield under stress was accounted for by translocation of stem reserves. Among the
117
cultivars, IR 64, IR 46 and Mahsuri had similar ATR values with no shade but IR 46
had more CHO coupled with high ATR and ACR under both shade and water deficit
conditions. Thus, IR 46 had the best CHO accumulation and translocation to confer
stress tolerance.
Table 10.
Variety
Treatments
Dry season
Yield
ATR
ACR
Yield
ATR
ACR
IR 46
No shade
Shade 10-25 DAPI
Shade 5-20 DAPE
4.0
3.1
2.8
5
10
24
20
21
35
3.9
3.1
2.8
13
22
34
20
23
40
IR 64
No shade
Shade 10-25 DAPI
Shade 5-20 DAPE
4.6
3.6
3.4
6
12
28
24
26
45
4.4
3.5
3.3
8
23
40
24
28
46
Mashuri
No shade
Shade 10-25 DAPI
Shade 5-20 DAPE
3.0
2.1
1.8
4
10
20
21
22
36
2.9
2.0
1.7
6
12
25
20
25
38
0.1
0.1
LSD 0.05
Reduction of rice grain yield by shade was also reported by Murty and Sahu,
1987; Venkateswarlu, 1977; Jadhav, 1987; Patro and Sahu, 1988). Yoshida (1973) found
that shade, particularly at reproductive and ripening phases reduced rice yields by
40 to 78 compared to unshaded controls. Additive effects of shade and water deficit
were observed in the dry but not in wet season trials. It was concluded that IR 46
had the greatest shade and drainage tolerance during panicle emergence and
flowering owing to its ability to form and set spikelets using accumulated assimilates
(Chaturvedi and Ingram, 1989).
CONCLUSION
118
Higher CHO at anthesis plus high ATR appear useful traits to select
cultivars for drought tolerance at flowering.
Yield was directly correlated with maximum ATR under stress condition.
IR46 had relataively more shade and drainage tolerance during flowering
stage owing to its ability to form and set spikelets using accumulated
assimilates.
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SUMMARY
Drought is a major problem for rice grown under rainfed lowland and upland
conditions, but progress in breeding for drought resistance has been slow. This paper
reviews genotypic variation in stress-related traits in rice, and suggests how stress
physiology can contribute to plant breeding programmes that aim to improve yield
under water-limiting environments.
There are several putative traits which confer drought resistance in rice. The
most important is the correct phenology which matches crop growth and
development to the water environment. A deep-root system and some other drought
avoidance mechanisms are useful in upland conditions. However, their usefulness
is not known for rainfed lowland rice where the development of a hard pan prevents
deep root penetration and where transpiration is a small proportion of total water
loss from the system. Drought tolerance mechanisms appear to offer more scope for
the genetic improvement of drought resistance in the rainfed lowland systems.
Osmotic adjustment is promising, as it can counteract the effects of a rapid decline
in leaf water potential and there is large genotypic variation for this trait. Green leaf
retention appears to be a useful character for a prolonged drought, but is affected
by plant size and this complicates the use of this as a selection criterion for drought
resistance.
A major reason for the slow progress in breeding for drought resistance in
rice is the complexity of the drought environment, which often results in the lack of
clear identification of the target environment(s). Then. is a need to identify the relative
importance of the three common drought types; early stress which often causes delay
in transplanting, mild-intermittent stress which can have a severe cumulative effect,
and late stress which affects particularly the late maturing varieties in rainfed lowland rice aerobic and anaerobic soil.
INTRODUCTION
Plant breeding aims to produce new cultivars suitable for a pre- defined target
populations of environments. Often the outcome is improved crop yield by
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strategies which are appropriate for the target population of environments. The focus
will be on rice, but work from other crops is included to make a comparison with
rice.
G X E INTERACTION AND SELECTION FOR DROUGHT RESISTANCE
Genotype by environment (GxE)interactions are expressed as a change in the
relative performance of genotypes in different environments. These interactions
complicate selection of superior genotypes when they are large relative to genotypic
variation and particularly when they result in a change in the rank of genotypes
across environments (Haldane 1946; Baker, 1988a). The incidence, size and nature
of GxE interactions are commonly investigated by conducting multi-environment
trials (METs), where a sample of genotypes is evaluated at a number of sites in a
number of years. Statistical models and methodology for the analysis and
interpretation of such series of experiments have been widely discussed in the
literature (Comstock and Moll, 1963; Baker, 1988b; Eisemann et al., 1990; Nyquist,
1991; Cooper et al., 1993; Cooper and De Lacy, 1994). Such analysis allows an
evaluation of the size of GxE interaction variation relative to genotypic variation for
the plant traits studied. The objective here is not to review GxE interactions in general
but to consider some aspects of how they can complicate interpretation and
extrapolation from the results of physiological experiments which concentrate on
genotype adaptation to water stress and/or genotype by water-stress (GxE)
interaction in rice grown under water limiting conditions. The two aspects which
will be considered here are (i) definition of targeted environments for the breeding
programme concerned and (ii) the relevance of particular types of physiological
experiments to the target population of environments. The salient principles are
developed by first considering a traditional statistical model used by plant breeders
to analyse the results of genotypic adaptation in METs. This is then re-defined in
terms of environmental factors which impinge on the results of drought resistance
research on rice.
Where genotypes are tested at a series of sites in a number of years, a linear
model which explicitly considers the cross- classification structure of sites and years
is often used to analyse the results of the experiments. This model may be formlly
defined as:
where Pijkl is observation 1 on genotype i at site j in year k and i=l, ..., ng, j=1,
..., ns, k=l, ... ny, l=1, ..., nr with ng , ns , ny and nr the number of genotypes, sites,
years and replicates, respectively; is the grand mean of all observations; gi is the
effect of genotype i; sj is the effect of site j; yk is the effect of year k; (sy)jk, (gs)ij,
(gy)ik, (gsy)ijk are the interaction effects associated with the main-effects; and e ijkl is
the random error effect 1 associated with genotype i at site j in year k. Generally data
on quantitative traits such as grain yield collected from METs are analysed in relation
to this model. From this model the relative importance of genotypic and GxE
interaction effects can be estimated for the traits analysed. Two deficiencies of such
an analysis are (i) that there is no definition of what aspects of the environment are
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water stress levels (refer to these by the symbol W for water-stress) are encountered
in different experiments. Other environmental factors such as temperature,
nutritional deficiencies and toxicities and biotic stresses (refer to these by the symbol
0 for other) will also influence the performance of the genotypes and these may
vary among experiments. Therefore, the model can be redefined as:
where Pijkl is observation l on genotype i under water-stress j and influenced
by other environmental factor k and i=l, ..., ng, j=1, ..., nw, k=1, ..., no , l=1, ..., nr with
ng, nw , no and nr the number of genotypes, water-stress environments, other
environmental factors and replicates, respectively; m is the grand mean of all
observations; gi is the effect of genotype i; w j is the effect of water-stress type j; ok
is the effect of other environmental factor k; (wo) jk , (gw)ij , (go)ik , (gwo)ijk are the
interaction effects associated with the main-effects; and eijkl is the random error effect
1 associated with genotype i under water- stress j and other environmental facror k.
Clearly this is a simplified model relative to that which would generally apply to
the target production-system. However, it applies to experimental situations where
we can evaluate genotype in a factorial combination of water stress regimes and
other environmental factors. This model is applicable to the sorts of physiological
experiments commonly conducted to investigate the physiological basis of drought
resistance such as our experimental types 1 and 2 above. An important difference
between models (1) and (2) is that developing and analysing series of experiments
in terms of model (2) asks specific questions about the nature of drought resistance
of genotypes in relation to types of water stress. However, to allow such an analysis
it is necessary to be able to identify types of water-stress environments and generate
these within the experiments. A model-based answer to the question of relevance
posed for experiment types 1 and 2 can be obtained by considering the experimental
scenarios posed in context with model (2). This is achieved by placing the appropriate
restrictions on the effects in the model and assessing how the components of model
(2) are confounded (Nyquist, 1991). The two experimental types are considered in
turn.
For experiment 1, where a range of water-stress environments is considered
but other environmental factors are held constant, no =1 and therefore k=1 in model
(2). Here estimates are obtained for genotypic effects across the water-stress regimes
and for GxW interaction. In this situation model (2) becomes:
In this presentation of the model the square brackets surround the effects of
model (2) which are confounded. In this case the estimates of the genotypic effects
are confounded with the genotype by other- environmental-factor (GxO) interactions
and the genotype by water-stress (GxW) interaction effects are confounded with the
genotype by water-stress by other-environmental-factor (GxWxO) interactions.
Therfore, as the GxO and GxWxO interactions increase within the target productionsystem, the capacity to predict the relative drought resistance of genotypes and
expression of GxW interactions in the target production-system from this type of
experiment will decrease. However, it should be emphasised that where GxW
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interactions are the major component of the GxE interaction complex and both GxO
and GxWxO interactions are small, the results from these types of experiments will
allow good prediction of the relative drought resistance of genotypes in the
environments of the target production-system.
For experiment 2, where only one type of water-stress environment is
considered and other environmental factors are held constant, n o =l and n w =l and
therefore j=k=l in model (2). Here estimates are obtained for genotypic effects only
and no interaction effects can be estimated. In this situation model (2) becomes:
In this case the estimates of the genotypic effects are confounded with all
three components of the GxE interaction complex. Therefore, in this type of
experiment there is no separation of the genotypic effects from the GxE interaction
complex and as any one of the GxE interaction components increases, the
predictive value of the specific environmental screen for the target
production-system will decrease. Again it is important to emphasise that while
prediction to many of the environments which comprise the target production
system may be unsuccessful, if the specific drought resistance screen matches
one type of target-environment in the production-system then the screen will
provide good prediction to that target environment. The additional question
which must be asked in this situation is what is the frequency of occurrence of
the particular target environment. For example, a seedling screen in a dry season
may be useful for identifying rice genotypes which perform relatively well under
early water stress conditions in a wet season, again assuming negligible GxO and
GxWxO interactions. If this form of water-stress occurs with high frequency in
the target production- system then the specific screen would be relevant.
However, if this environment only occurs at a low frequency then while the screen
may have good predictive capacity, its relevance would be decreased.
From the above consideration of screening genotypes for drought resistance
it is argued that a critical component of studying drought resistance is a clear
definition of the relevant target environments within the crop-production system.
Where the objective is to select genotypes or identify physiological components
which contribute to drought resistance, consideration must be given to the relative
contribution of GxW, GxO, and GxWxO interactions in relation to the target
environments. Without due consideration of the relevance of types of experiment,
the success of a particular drought resistance screen, putative drought resistance
trait or plant ideotype is likely to be poor in its contribution to plant breeding
strategies and progress in breeding for drought resistance will continue to be slow.
At present little is known of the size and causes of GxE interactions in rice production
under water limitation. Given the ubiquitous nature of GxE interactions in other
crop production systems (Delacy et al., 1990) and the diverse nature of the types of
environments in which rice is grown it may be anticipated that GxW, GxO and
GxWxO interactions will be large for quantitative traits such as grain yield. In the
following sections we consider putative drought resistance traits which have been
studied in rice and assess their possible contribution to drought resistance for various
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types of drought conditions. Specific attention is given to the incidence and influence
of GxE interactions on the role of these putative traits in drought resistance.
THE PHYSIOLOGICAL BASIS OF INTERACTING FACTORS
There are a number of reasons for the genotype by drought environment
interactions discussed above. One reason is that yield potential under favourable
water conditions varies among genotypes but yield under water limiting conditions
may be similar. This interaction is not of interest here as we try to understand yield
differences in water limiting conditions.
Timing of drought development in relation to a genotypes phenology is
important for GxE interaction. It is well known that panicle development to
anthesis is the most critical stage for water stress in rice. Our work (Boonjung et
al., 1994b) with rice indicates that grain yield is lowered at the rate of 2% per day
as a 15-day stress period (dawn leaf water potential less than -1.0 MPa) later
during panicle development is delayed (Fig. 1). Unfilled grain number increases
sharply with stress during late panicle development. Water stress reduces
assimilate production and this causes reduction in yield components such as
panicle number per plant, spikelets per panicle, filled grain percentage, and
individual grain mass. However, unfilled grain number appears most susceptible
to reduced assimilate availability.
Assuming the reduction of 2% grain yield per day with the delay in
termination of 15 days stress, a 20-day difference in flowering time between two
cultivars would cause a grain yield difference of about 40% if they have the same
yield potential under non-limiting conditions. Thus, it is likely that the cultivars
with a difference in phenology will react differently to a drought, depending on the
timing of stress development (Maurya and OToole, 1986). Phenology is important
in determining grain yield response also because quick maturing cultivars often
escape from severe stress, while late maturing cultivars may be affected by a terminal
stress. These results suggest that genotypes should be compared for their drought
resistance / susceptibility within the same phenology group, or at least genotypic
variation in phenology should be corrected statistically before genotypic difference
in drought tolerance is estimated (Bidinger et al., 1987a, b). Alternatively it is possible
in some experiments to implement a strategy of staggered planting of different
varieties so that they would flower at about the same time (Lilley and Fukai, 1994b).
In addition to timing of drought in relation to crop phenology, duration and
severity of drought can also affect genotypic response to drought. For example,
genotypic responses to mild, intermittent stress may be quite different from those
to prolonged stress.
Another reason for GxE interactions among drought environments is that
drought affects crop growth indirectly through contributing to the incidence of
another adverse condition. For example, certain diseases in rice tend to develop
under dry conditions. Dry conditions may not always induce the disease in the same
manner. Thus, where there is genotypic variation in resistance to the disease in some
dry years, some lines appears to be doing well relative to others but not in other
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Fig. 1.
Relationship between grain yield under water stress conditions, relative to the corresponding
sowing in the irrigated trial, and the time of water stress termination expressed in days to
anthesis, of three trials. Experiment 1 (0, Exp 1), Severe stress
SS) and mild stress
(
MS) trials in Experiment 2. S1- S4 indicate time of sowing. (Boonjung et al 1994b).
dry years, depending on the pattern of disease development. Under these situations
the specific drought resistance response is confounded with the presence of the other
factor, a disease in this case, and the large GxWxO interactions (see equations (3a)
and (3b) complicate interpretation of drought resistance responses. It is then
necessary to identify the presence of the disease in the experiment and screening
methods for drought resistance need to consider the complications generated by this
other factor. Where the disease occurs widely with drought, the direct selection
against the disease may be more appropriate than attempting to develop cultivars
with drought resistance per se.
Rainfed lowland rice has a unique nature of alternating aerobic and anaerobic
soil conditions during the crop growth cycle and this is likely to be another major
source of the GxE interactions, particularly GxO and GxWxO interactions. During
a flooded period, rainfed lowland rice may perform similarly to irrigated lowland
rice, but often growth is reduced as water level recedes to below the soil level. Our
recent experience in Northeast Thailand indicates that the rice plants do not show
typical water stress symptoms of leaf rolling and leaf death immediately after the
flood water has disappeared from the soil surface. In some cases these symptoms
were never seen, but grain yield was generally low in the sites where there was no
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flooding water during the latter part of the growing season. This may be related to
phosphorus becoming less availabe in the aerobic soil, or to a loss of nitrogen during
alternating periods of aerobic and anaerobic soil conditions (Patrick and Wyatt, 1964).
It is also possible that loss of standing water results in increased acidity of soil water,
as in many cases the soil is strongly acidic. With standing water the acid soil is
unlikely to cause a problem. Rice is known to be susceptible to manganese and
aluminium toxicity and reduced pH of the soil solution may induce the toxicity
within a short time period.
Genotypes may differ in their response to the manganese or aluminium level
in the soil as found in some crops. If experiments are conducted at different locations
with soils of various pH levels, there could be strong GxE interactions in dry seasons
and between wet and dry years.
Where the objectives is to study drought resistance, the incidence of the above
complex mixture of these and other types of environment in rice production systems
and their potential for generating GxE interactions strongly suggests (i)the
importance of identifying the types of stress environments sampled in the traditional
METs conducted in plant breeding programme and (ii) more focussed screening of
lines under relevant target environment conditions. In both of the areas there is scope
for stress physiology research to contribute to plant breeding strategies.
PUTATIVE TRAITS FOR IMPROVING DROUGHT RESISTANCE
This section evaluates the evidence for a number of putative traits which may
confer drought resistance in rice. The use of putative traits to improve grain yield
is a classic example of indirect selection in plant breeding. In this case the objective
is to indirectly increase grain yield, or stability of grain yield by selecting for putative
drought resistance traits, or for a combination of yield and the traits. The indirect
response to selection( Gy/x) for character y (yield) from selection for character
x(putative draught resistance trait) is described by the equation :
where ix is the standardised selection differential applied to the putative
drought resistance trait x; hx and hy are the square root of the heritability of the
putative traits x and y, respectively; rgxy is the genetic correlation between traits x
and y; and py is the square root of the phenotypic variation for character y. This
equation may be expressed in different forms and a detailed treatment of the
principles of indirect selection can be found in Falconer (1989) and in the review by
Gallais (1984). The component of equation (4) which determines whether there is
scope for indirect improvement of yield via the putative trait (s) is the genetic
correlation between the two traits. Genetic correlations are a result of either linkage
or pleiotropy. For a high chance of successful indirect selection, identification of a
putative trait which is pleiotropic for yield is desirable. Linkage relationships can
change with time and between populations due to recombination. Recently Aastveit
and Aastveit (1993) considered the impact of GxE interactions on genetic correlations
between traits. In general, the presence of unpredictable GxE interactions will reduce
the heritability of traits and the reliability of genetic correlations between traits
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Root system
Under upland condition, rice, maize and sorghum have similar root length,
density and water extraction pattern down to 60 cm depth (Fukai and Inthapan,
1988). However, below that depth, water extraction by rice is very small and this is
the main reason for the susceptibility of rice to water strees, compared with sorghum
(Inthapan and Fukai, 1988). The develoment of the rice root system is also sensititive
to soil water deficit (Boonjung et al., 1994a).
O, Toole (1982) suggested three root-related adaptive mechanisms for rice. (1)
For relatively large soil water reservoir, increase in rooting depth, root density,
root-shoot ratio and possibly also root conductance. For relatively small soil water
reservoir, (2) possibly increased root penetration of any physical-chemical
impediment and (3) possibly osmotic adjustment of root system. High osmotic
adjustment of roots would allow more thorough extraction of soil water and also
enhanced dehydration tolerance which would enhance root system survival and
plant recovery upon rewatering. However, experimental results to date are not
available to show the effects to osmotic adjustment in roots. Most work on the root
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system of rice in relation to drought resistance has been conducted under upland
conditions where there is a large soil water reservoir. This relates to OTooles first
point of the adaptive mechanisms, and emphasises the importance of rooting depth,
mot density, and mot thickness.
The development and function of the rice root system, and their relation to
drought resistance were reviwed by Yoshida and Hasegawa (1982). Their review
was mostly based on upland work conducted at IRRI in the 1970s, and their major
findings are summarised here. Rice has a well developed mot system near the soil
surface (0-30 cm), but generally not below 30 cm, compared with other major crops.
There is, however, a lagre variation in root length density below 30 cm among rice
genotypes. Generally genotypes with high root length density below 30 cm have a
deeper root system. Yoshida and Hasegawa used the ratio of deep root weight and
shoot weight as an index for drought resistance (avoidance) as the large deep mot
system would be able to extract more water and the small shoot would require less
water. They found the ratio to be positively related to field evaluation of drought
resistance. Using 1081 entries, they also found that the entries with large deep
root-to-shoot ratios tended to be taller and have less tillers. These characters match
the description of traditional upland rice varieties, since the rice root system is
composed basically of nodal roots, genotypes with large tillers tend to have more
roots, particularly on secondary and tertiary tillers which appear late and have short
mots. Thus, it appears that genotypes with a small number of well-developed tillers
have a smaller number of longer roots and this results in high root length density
at depth. They would be thick (larger diameter) and hence, have high conductance
with large xylem vessels.
While a direct relationship between root length at depth and the amount of
water extracted from the layer is rare. Puckridge and O, Toole (1981) found a deep
rooting cultivar, Kinandang Patong, that extracted a larger amount of water at 40-70
cm depth than the two cultivars IR20 and IR36 which were shallow rooted. Similar
results were shown by Mambani and Lal (1983b, c). Our recent work, also under
upland conditions, (Lilley and Fukai, 1994a) shows that variation in water extraction
among four cultivars was directly related to the variation in root length density.
Thus when the amount of water extracted from a 20 cm layer in the soil profile is
plotted against the root length density in the layer, the data from the four cultivars
fall on one curve for each experiment (Fig. 2). Incresed root length density promoted
the rate of water extraction, though the duration of linear phase of water extraction
was to some extent reduced. In their experiments, ground cover varied among
cultivars, but that apparently did not affect the root length density-extracted water
relationship, indicating the importance of mot length in determining the amount of
water extracted.
Lines with high root length density tend to have high leaf water potential and
delayed leaf death due to water stress (Mambani and Lal, 1983a; Cruz and OToole,
1985; Ekanayake et al., 1985a). These favourable conditions may result in higher grain
yield in lines with high mot length under water limiting conditions(Mambani and
Lal, 1983a), although a yield advantage is not always observed (Puckridge and
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Fig. 2.
Relationship between a) extractable water, b) maximum extraction rate and c) effective duration
of extraction and root length density in a 0.2 m layer (0.2-0.8m depth) for four cultivars
( CPICB, Lemont, Rikuto Norin 12, Todoroki Wase) in the vegetative stage (solid
symbols) and reproductive stage (hollow) trials. (Lilley and Fukai, 1994a).
OToole, 1981). In the case of Lilley and Fukai(1994b) mentioned earlier, there was
an indication that the cultivar with the highest root length performed better than
others under mild stress conditions, but there was no direct relationship between
total root length and grain yield in the experiment where there was only one period
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of drought. It is likely that the advantages of varieties with larger root systems would
be greater when there are several drougth periods, i.e. intermittent stress.
However, it is unlikely that the genotypes with high mot length density at
40-80 cm depth will have any advantage under most rainfed lowland conditions, as
the hard pan which develops closer to the soil surface would preclude the expression
of the potentially superior root system. It is rather the genotypes ability to penetrate
the hard pan that may be more important in rainfed lowland conditions. In a lowland
experiment where water stress developed from 45 to 75 days after transplanting,
grain yield of 30 genotypes was not related to any mot characters determined in
aeroponically grown plants (Ingram et al., 1990). Nevertheless in some rainfed
lowland fields where puddling is not practiced, for example due to sandy soil texture,
high root length at depth may have direct relevance.
Root axial resistance can be important in determining the rate of water flow
to the shoot. Rice plants are often not able to extract water thoroughly from deeper
layers, partly because of low root length density at depth, but also increased axial
resistance due to the increased distance to the shoot and also to the smaller diameter
of the roots. Genotypic variation in mot thickness is associated with that in xylem
size (Yambao et al., 1992). However, they consider that increased xylem vessel
diameter will not directly increase the drought resistance of a genotype.
Determination of root length and root depth in the field is time consuming,
and quicker methods have been devised which could be used in screening for an
ideal root system in a breeding programme. For example aeroponic and hydroponic
culture can be used for root system determination (Loresto et al., 1983; Ekanayake
et al., 1985 a; Haque et al., 1989). However, the value of this strategy is questionable
for rainfed lowland rice given the findings of Ingram et al. (1990). Gomathinayagam
et al. (1989) suggest that seminal root growth can bo used for screening deep rooted
rice. In the field the root pulling resistance can be used as a measure of root systems
(O, Toole and Soemenatono, 1981). Plants with high mot pulling resistance were able
to maintain high leaf water potential (Ekanayake et al., 1985b).
Some root system characteristics appear to have high heritability. Using F1,
F2 and F3 populations of the cross between IR20 (a shallow, thin root system) and
MGL-2 (a deep, thick root system), Ekanayake et al. (1985a) found high heritability
for root thickness, mot dry weight and root length density. However, heritability for
root pulling resistance was rather low (Ekanayake et al., 1985b).
Shoot-related mechanisms
OToole (1982) suggested three shoot-related adaptive mechanisms;
accumulation of amino acids or growth regulators, drought avoidance mechanisms
and osmotic adjustment.
Growth regulators
Little work has been conducted with rice in relation to growth regulators and
drought resistance. Dingkuhn et al. (1991a), in the same experiment as will be
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reported later for osmotic adjustment and other physiological characte of seven
diverse cultivars by Turner et al. (1986a, b), found cultivar differences in abscisic acid
(ABA) accumulation, but the difference was not related to physiological behaviour
of cultivars under water stress. On the other hand, they found stress-induced proline
accumulation to be positively correlated with osmotic adjustment.
Avoidance mechanism
In contrast to a deep root system which can increase water uptake, avoidance
mechanisms in the shoot reduce water use, thus avoiding the development of low
plant water potential.
Epicuticular wax is an avoidance mechanism trait which may have a
significant effect on rice growth during water stress and recovery upon rewatering.
However, the contribution of this putative trait has not been examined in details.
Rice is known to have much less epicuticular wax than other cereals and it has
generally high cuticular (epidermal) conductance (O'Toole et al., 1979). This implies
that rice will lose water even when stomata are completely closed, and this may
lead to rapid leaf death. There is genotypic variation in quantity of epicuticular wax
(OToole, 1982). In our recent experiment, epidermal conductance was found to vary
among rice cultivars, but there was no indication that cultivars with low epidermal
conductance retain green leaves for a longer period.
Leaf diffusive conductance and leaf rolling are also avoidance mechanisms,
for which genotypic variation is well documented (O'Toole and Cruz, 1980). The
variation is at least partly due to differences in the ability to extract water from the
soil, as a result of differences in root system. This causes differences in leaf water
status (water potential), and genotypes which maintain high leaf water potential
tend to maintain high leaf conductance and low leaf rolling(OToo1e and Moya, 1978).
However, the leaf rolling score and leaf water potential relationship varies among
cultivars (Turner et al., 1986 a; Fukai and Inthapan, 1988) possibly because of cultivar
differences in osmotic adjustment. Cultivars with high osmotic adjustment can
maintain turgor potential at relatively high levels under low leaf water potential.
This relationship has been documented in other cereals, though strong experimental
evidence is lacking in rice. Within a genotype, osmotic adjustment is known to affect
leaf rolling in rice (Hsiao et al., 1984).
The direct effect of these drought avoidance mechanisms on growth of rice
plant under limited soil water conditions is not certain, particularly in rainfed
lowland conditions where the amount of water used as transpiration would be a,
rather, small proportion of total water use. The latter includes evaporation from
water or soil surface, deep percolation and seepage. While Dingkuhn et al. (1989a)
found under mild stress conditions a correlation between leaf rolling and water
potential, suggesting that leaf rolling has a positive effect in maintaining high leaf
water potential, our experience is that cultivars which are able to maintain high leaf
water potential have a small degree of leaf rolling. This indicates that, generally leaf
rolling is a result of other avoidance mechanisms, which result in high leaf water
potential.
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Osmotic adjustment
Osmotic adjustment is an adaptive process in which solutes accumulate in
the cell and osmotic potential is decreased in response to adverse growing conditions
such as drought, salinity and low temperature. As water stress develops osmotic
adjustment is induced, and as such osmotic adjustment increases with decrease in
leaf water potential (Steponkus et al., 1982; Turner et al., 1986 b). Development of
osmotic adjustment however, appears to be rapid in rice, perhaps because of rapid
development of water stress as measured by leaf water potential, compared to
sorghum and maize (Fukai and Inthapan, 1988). Thus any benefits of high adjustment
would be expected in rather early stages of a drought period in rice.
With osmotic adjustment, turgor pressure is maintained at a relatively high
level in spite of reductions in leaf water potential (Cutler et al., 1980). Development
of leaf rolling and half death can be delayed by osmotic adjustment (Hsiao et al.,
1984). It should be pointed out however, that turgor is not necessarily the sole factor
determining the responses of these processes to soil water deficit.
The early work by Steponkus et al. (1982) using four cultivars has shown rather
small variation in osmotic adjustment in rice. The maximum adjustment was about
0.3-0.5 MPa. In the field studies, there was some variation in osmotic adjustment
with a maximum of 0.5 MPa in lowland cultivars and less in upland cultivars among
seven diverse cultivars of rice (Turner el al., 1986b). However, the differences were
related to the development patterns of water stressed cultivars, and the greater
degree of apparent adjustment in the lowland cultivars was due to the greater degree
of cumulative stress. Yang et al. (1983) found some difference in the osmotic potential
water potential relationship between two constrating cultivars, Taichung Native 1
and OS4.
There is no evidance to indicate that genotypic variation in osmotic adjustment
has a positive effect on growth and grain yield in rice, though probably there has
been no serious attempt to test the usefulness of this character in the field. Our recent
work however, does indicate some positive effect of osmotic adjustment in
counteracting low leaf water potential to provide better green leaf area retention
(Henderson et al., 1993). Because osmotic adjustment in rice reaches its maximum
value quickly and the maximum value is maintained for a while, it may be effective
in buffering against the deleterious effect of water stress under situations of rather
a small soil water deficit as may occur under intermittent stress.
The mechanisms of the effect of genotypic variation on yield are well
documented in sorghum, and are briefly reviewed here. Several field experiments
using 2-6 commercial hybrids with a contrast in osmotic adjustment have shown
clear yield advantages of hybrids with high osmotic adjustment when water stress
develops before anthesis or during grain filling (Wright et al., 1983, Ludlow et al.,
1990, Santamaria et nl., 1990). Grain yield under stress conditions relative to that
under well-watered conditions increased linearly with the extent of maximum
osmotic adjustment of the line. Compared to genotypes with low osmotic
adjustment, those with high osmotic adjustment were able to extract water more
thoroughly, maintain higher grain number and translocate a greater amount of
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pre-anthesis assimilates to fill grains. Using lines from a sorghum population that
had differences in osmotic adjustment, Tangpremsri (1989) showed a yield advantage
of about 0.5 t ha1 in the high osmotic adjustment group. These groups had similar
yield under well watered conditions. Under water-limiting conditions, the high
osmotic adjustment group produced larger maximum leaf area, better leaf retention
during grain filling and more grains per plant. Thus, there are a number of
experimental results to suggest the advantge of high osmotic adjustment when
sorghum experiences a prolonged period of water stress.
The current work by J.M. Lilley and M.M. Ludlow at CSIRO, Brisbane,
Australia, shows a larger genotypic variation in osmotic adjustment than previously
observed in rice. Measurements on rice plants grown in pots in a controlled
environment and subjected to slow (25 day) drying cycle showed that there is a large
range in maximum osmotic adjustment (0.4 - 1.7 MPa) among a set of 59 lines of
diverse adaptation/background. Average osmotic adjustment of lines from upland
environments was lower (0.73 MPa) than that of rainfed lowland, irrigated lowland
and deep water ecotypes (0.96-1.03 MPa). However, there was a large range in
osmotic adjustment within each ecotype.
They also examined variation for osmotic adjustment among a sample of
recombinant inbred lines from a biparental cross. The parents, Morobereken and
CO39, had low (0.7 MPa) and moderate (1.4 MPa) osmotic adjustment, respectively.
Osmotic adjustment of the 52 recombinant inbred lines (RIL-F7), which had been
mapped with RFLP markers, ranged from 0.7-2.7 MPa (Fig. 3). The frequency
distribution of level of asmotic adjustment in the RILs was continuous and there
was no evidence for osmotic adjustment being simply inherited. Transgressive
segregation was observed with 6 lines having significantly higher osmotic
adjustment than the high parent CO39. Morgan (1991) reported that a single recessive
gene accounted for the variation in osmoregulation observed in wheat. Our work
in sorghum (Basnayake et al., 1994a, b) suggests that the inheritance of osmotic
adjustment is at least oligogenic. However, two major genes for high osmotic
adjustment were identified. Current work is investigating the contribution of these
genes to yield under water stress conditions.
The large genotype variation for osmotic adjustment in rice offers considerable
scope for improving these traits in current cultivars. Screening for osmotic
adjustment currently requires intensive measurement procedures and is time
consuming and expensive. Molecular markers such as RFLPs and RAPDs could be
identified for osmotic adjustment genes. The variation in the populations offers a
good possibility of obtaining suitable RFLP markers for these traits so that these
traits could be readily incorporatd in rice breeding programmes.
Water use efficiency
Water use efficiency (WUE) is commonly defined as dry matter production
per unit of water used, and is closly related to transpiration efficiency, dry matter
growth per unit of water transpired. Instantaneous measurements of leaf
photosynthesis and transpiration used to approximate WUE or transpiration
138
Fig. 3.
efficiency. Using this technique Dingkuhn et al. (1989b) found WUE to be high in
tropical japonica types, intermediate in indica types and low in aus types, and also
considerable variation in WUE within the types. On the other hand Fukai et al. (1985)
found no significant variation in WUE among four temperate japonica cultivars. They
however, found WUE to decline rapidly with leaf ageing. One of the difficulties of
estimating WUE from instantaneous gas exchanges is the fluctuation of WUE, due
to leaf age or growing conditions such as vapour pressure deficit. This problem can
be overcome by use of stable carbon isotope analysis. Dingkuhn et al., (1991b) found
a good relationship between WUE determined by the gas exchange method and
carbon isotope discrimination. In contrast to drought avoidance mechanisms
expressed in the shoot which decrease the rate of assimilate production, there is no
cost for high WUE, and hence cultivars with high WUE would be advantageous in
dry conditions, unless WUE is associated with characters which are undesirable for
high yield.
Green leaf retention
Since leaf death occurs as a result of severe drought and is an easily
recognisable character in the field, green leaf retention is often used as a selection
139
criterion for drought resistance in rice (De Datta et al., 1988), assuming that cultivars
with good leaf retention produce a higher yield than others in dry environments. In
rice, 30 days old seedlings are subjected to drought and they are visually scored
using a standard (drought score) developed by IRRI. Lines which have better green
leaf retention also recover better after water stress is relieved (Malabuyoc et al., 1985).
Drought at the seedling stage is commonly related to leaf water potential
(Chang et al., 1979). Lines which can maintain high leaf water potential tend to retain
green leaves and hence, have a low drought score. Thus, low drought score generally
indicates that the lines possess mechanisms of drought avoidance, although some
lines appear to be able to maintain green leaves better than others at low leaf water
potential and hence, they are dehydration tolerant (Henderson et al., 1993). Our
recent experiments indicate that genotypic variation in drought score is strongly
associated with variation in light interception prior to the commencement of a stress
period. Thus, lines with large plant size, and hence, with high light interception were
stressed more quickly and had high drought scores. Variation in drought score of a
line is also associated with variation in plant size obtained under various water stress
environments, i.e for some reasons a line may establish more quickly than others in
one site but not in others and this causes inconsistency in relative drought score
among lines. When full ground cover is achieved before drought development, the
genotypic variation in plant size becomes less important in determining leaf death
and this may be a reason for change of genotypic ranking on drought score between
young seedlings and plants with a well developed canopy (Henderson et al., 1993).
In a rainfed lowland experiment where drought was developed in a crop at a high
plant density from 45 to 75 days after transplanting, genotypic variation in drought
score of 30 lines was found to be correlated with yield (r=-0.66) (Ingram et al., 1990).
While leaf death occurs as a result of drought, it does not occur until plants
are severely stressed. Under upland conditions, Boonjung et al. (1994c) have shown
that leaf death is least sensitive to soil water deficit and leaf elongation and leaf
diffusive conductance are much more sensitive. This implies that in rather mild and
intermittent stress where available soil water does not decrease below 50% of the
total available water, plant growth will be affected because of reduced leaf expansion
growth and photosynthesis. Genotypic variation in these characters, rather than
drought score, would determine crop growth. In one of our upland rice experiments
we found no association between leaf conductance and drought score among 20
lines examined, indicating that selection for low drought score may not necessarily
select for high leaf conductance under stress conditions.
Lines with low drought scores do not necessarily perform well in dry
conditions (Puckridge and O Toole, 1981). This again shows the importance of
identifying the target environment and developing an adequate screening method
suitable for the environment. The use of drought score at the seedling stage may be
appropriate for drought that develops early in the season in rainfed lowland; for
example in July in Thailand, particularly for direct seeded rice.
One advantage of lines with good green leaf retention is that they tend to
recover quickly when the stress period terminates. This may be particularly
140
important when stress develops around panicle initiation, as the lines with good
leaf retention can supply more assimilates to the developing panicle. This in turn
will result in production of a large number of spikelets (Lilley and Fukai, 1994b).
Production and retention of fertile spikelets
When a rice variety is grown under different environments, grain yield is
mostly related to grain number as single grain weight is stable across environments.
Grain yield is mostly limited by grain sink capacity to accept assimilates to fill grains
(Fukai et al., 1991). Grain number is determined by the number of spikelets at anthesis
and proportion of spikelets which produce filled grains (filled grain percentage).
The number of spikelets is directly related to the rate of assimilate production
between panicle initiation and anthesis, regardless of whether the assimilate
production is altered by water stress or shading (Boonjung, 1993). Filled grain
percentage on the other hand is related to assimilates produced around anthesis,
and is particularly susceptible to water stress (Cruz and O Toole, 1984, Boonjung,
1993). Maurya and OToole, (1986) found large genotypic variation in filled grain
percentage during the dry season. Reduction in leaf water potential at anthesis may
cause poor exertion of the panicle (Cruz and OToole, 1984; Ekanayake et al., 1989)
and increase the percentage of sterile spikelets because of pollination abnormalities
(Ekanayake et al., 1989, 1990). Ekanayake et al. (1989, 1990) showed some differences
in flowering response to water stress between the two cultivars they studied, and
their work suggests that drought resistant varieties should be able to maintain high
leaf water potential at anthesis and do not show pollination abnormalities as water
potential declines, compared to susceptible varieties.
of occurrence of these three types of water stress environments in the rainfed lowland
regions needs to be determined. Alternatively, if the components of water balance,
particularly seepage and deep percolation, are known for the paddy, a water balance
model may be used to estimate the level of perched water table during crop growth
and this could be used as an indicator of likely plant water stress. Comparison of
grain yield under rainfed conditions and irrigated conditions will provide
information on the magnitude of the drought problem.
It is likely that appropriate traits can be selected once the target environment
is accurately defined. Soil factors are also important in selecting traits, as mentioned
earlier for the root growth, acidity and nutritional problems. Lowered perched water
table in a dry period could interact with these factors, and affect growth indirectly.
Identification of these interactions should help in determining which traits are to be
considered in a plant improvement programme.
The following is a list of suggested putative traits for enhancing the drought
resistance of rice under different water stress conditions. It is emphasized that the
usefulness of each trait for conferring high yield under drought conditions is
unknown in most cases, therefore the traits listed, particularly for lowland
conditions, should be considered as those on which concentrated research effort may
be made by physiologists and plant breeders.
For different cultural conditions
Upland conditions:
Appropriate phenology, deep root system, shoot drought avoidance mechanisms.
Lowland conditions with hard pan:
Appropriate phenology, green leaf retention, tolerance for adverse soil conditions,
high root conductance, root penetration ability.
Lowland conditions without hard pan:
Appropriate phenology, tolerance for adverse soil conditions, deep mot system.
For different types of stress
Early season stress:
Transplanting tolerance, green leaf retention, recovery growth.
Mid season, intermittent stress:
Avoidance mechanisms, appropriate phenology, spikelet retention.
Late season stress:
Appropriate phenology, green leaf retention, tolerance for adverse soil conditions.
Recently crop models have been considered as possible tools to assist
investigation of the physiological and genetic basis of genotype adaptation (Shorter
et al., 1991) and selection decisions in plant breeding (Muchow et al., 1991). Currently
models are not at the level of development where they can be applied to the large
number of unknown genotypes evaluated in breeding programmes. However, they
may be useful in characterising the frequency of the early, mid-intermittent and late
stress environment types in the target population of environments. Our current
143
studies will evaluate the use of a rice model in this role in the rainfed lowland system
in Thailand.
CONCLUSION
Crop physiology can make a strong contribution to a plant breeding
programme designed for improving drought resistance. At present any
improvements in drought resistance are largely a fortuitous outcome of the empirical
multi-environment testing strategies used in the breeding programmes. While this
traditional approach has been effective, its efficiency is widely considered to be low.
We have identified the incidence of strong genotype by water stress (GxW)
interactions for putative drought resistance traits and grain yield as a major
constraint to the genetic improvement of drought resistance. Therefore, plant
breeding strategies and physiological research focussing on drought resistance in
rice must be conducted in a way which allows assessment of the effects of GxW
interactions. We consider that this can be achieved by more effective characterisation
of the type of stress encountered in individual experiments and relating this to
relevant target environments in the rainfed lowland production system. If this
approach is adopted, putative drought resistance traits can be evaluated in context
with the components of indirect response to selection in a way which accommodates
the effects of GxW interactions.
A number of putative drought resistance traits have been identified. The most
important for all drought environments is the correct phenology which matches crop
development to the water availability. Nevertheless, intermittent stress may develop
at flowering time, or the rainy season may finish earlier in some years and the crop
is subjected to stress at flowering. Genotypic variation in sensitivity of flowering to
stress is worth further investigation.
The importance of a deep root system is established as a requirement for rice
cultivars suitable for upland conditions as it ensures greater extraction of water held
in deeper soil profile and therefore, maintenance of a high leaf water potential during
a drought period. However, this is unlikely to be the case for rainfed lowland
conditions, where the existence of hard pan inhibits root penetration to deeper layers.
Water conservation strategies may also have limited success in rainfed lowland
conditions, because of the small quantity of water that can be stored in the soil
compared to the amount of water loss due to deep percolation, seepage and
evaporation from the lowland field. For both upland and rainfed lowland conditions,
cultivars with thick roots are advantageous in maintaining high water flow and
hence, favourable plant water status.
It has been shown that some rice lines have better dehydration tolerance than
others, judging by good green leaf retention even at rather low leaf water potential.
This appears particularly useful for rainfed lowland conditions where probably a
water- stress avoidance strategy has limited value. Dehydration tolerance may be
achieved by increased osmotic adjustment, but this needs further testing. Osmotic
adjustment has been found to vary greatly among rice genotypes.
144
Fig. 4.
A flow diagram showing processes of the effect of drought on grain yield, and important
factors associated with each process.
It appears also that soil water deficit in rainfed lowlands induces changes in
the environment which are not directly associated with plant water stress but may
affect growth greatly. Soil conditions change from anaerobic to aerobic during cycles
of flooding and drying, and there would be associated changes in nutrient availability
or toxicity. Genotypic variation in response to these changes appears worth further
investigation. Compared with the research effort for drought resistance in upland
rice environments, the research input to the rainfed lowland conditions has been
minimal.
A high priority research area is an accurate definition of the target population
of environments, particularly patterns of drought development. This would assist
definition of optimal multi-envimnment testing strategies for a breeding programme,
ensure determination of the best phenology group for the target environment, and
also focus areas of physiological research which can be directly related to the
objectives of a plant breeding programme.
145
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SUMMARY
The Rainfed Lowland Rice Research Consortium site in Bangladesh is located
at northwest part, known as Barind Tract, which is characterized by low annual
rainfall. Rice crop in this region often faces drought stress at reproductive phase.
Soil is poor in organic matter with hard and thick (8.0 cm) ploughpan. Two hundred
and fortyeight advanced breeding lines and varieties were screened and evaluated
with respect to different plant characters under field condition to find out suitable
varieties which can tolerate drought at later growth stages. Fortytwo advanced
breeding lines were identified as drought tolerant, out of which two
lines-BR4974-45-9-2 and BR4974-42-1-3 have deep rooting ability and can even
penetrate such hard ploughpan and also produce higher grain yield than the local
varieties under rainfed condition. Another line, IR41054-81-2-3-22 identified as early
maturing was found to escape drought and produce about 30% more grain yield
than the local popular early maturing variety. Soil management manipulation
treatment for reducing the ploughpan thickness through deep tillage and ploughpan
perforation through dhaincha (Sesbania aculeata ) roots significantly increased grain
yield and root mass of rice genotypes.
INTRODUCTION
About 36% of South and Southeast Asian rice growing areas come under
rainfed lowland rice ecosystem (BRRI, 1992). Uncertainty characterizes rice farming
in this ecosystem where it often suffers from drought, floods, pests, weeds and soil
constraints, sometimes all in the same field and in the same season. Rainfall is erratic
and other conditions for growing rice crop are diverse and unpredictable.
In Bangladesh, about 44% of total rice area is occupied by rainfed lowland
rice (transplant aman), out of which about 50% is covered by high yielding varieties
(HYV) developed by Bangladesh Rice Research Institute (BBS, 1992). Rest of the
areas are still being covered with traditional varieties, characterized by low yield
and photoperiod sensitivity enabling for late planting due to late recession of flood
water and late harvesting of preceding aus and jute crops.
The existing HYV are not suitable for cultivation under unfavourable rainfed
lowland situation and may suffer from drought, low temperature and soil stress at
1 Bangladesh Rice Research Institute, Gazipur, Bangladesh
reproductive and ripening phases particularly when the crop is late planted.
Therefore, steps are to be taken to develop varieties and associated technologies to
overcome drought, low temperature and soil stresses.
Drought in rice is a real concern and a serious yield limiting factor on much
of the worlds 70 million hectares of non- irrigated rice (OToole and Chang, 1978).
Lack of rainfall and the resultant soil moisture deficit form the basis of drought. TO
overcome drought stresses, plants develop some defensive mechanism such as
stomatal closure, thick cuticle, leaf rolling, profuse, or deep and thick root system
(OToole and Chang, 1978). Early maturity is also a way to escape drought stresses.
Extensive study is difficult to test all the above characters for a large number
of varieties/lines. Field screening by visual scoring would appear to be an indirect
means of screening for root system development (OToole and Chang, 1978). The
percentage of filled grain on the panicle is a useful criteria in ascertaining reactions
to reproductive phase drought. Proportional reduction in yield in a genotype due
to stress acts as a indicator of drought resistance or susceptibilitiy (OToole and
Chang, 1978). Plant height and panicle exertion is also an indication to resistance or
susceptibility to drought. Root system plays an important role under water deficit
conditions and the nature and extent of root development are major factors
governing plants response to moisture condition (Russell, 1959). Haque et al.,
(1988) also stated that selection based on root characters for drought resistance could
be done in the early stage of growth. A thick and dense ploughpan layer restricts
the rootzone volume of soil and hence reduces its moisture storage capacity.
The rainfed lowland rice research consortium site located in the Barind Tract
has a ploughpan at shallow depth (about 10 cm) which creates adverse conditions
for root development of rice crop. The Barind tract receives less annual rainfall than
most of the parts of Bangladesh (Fig. 1).
The distribution of rainfall is concentrated in a few months, from June to
September (Fig.2).
Transplant aman often faces terminal drought stress. The soil in this area is
also low in organic matter content, which adversely affects other soil properties
(Table 1). Rainfall is the only source of water for crop growth because large scale
irrigation is not possible due to undulating land scape and poor aquifers. To address
some of these problems a number of experiments were conducted at the site from
1991 to 1993.
Experiment 1. Screening of advanced breeding lines for drought tolerance at
reproductive phase under rainfed condition.
151
Fig. 1.
5m x 2 rows plot with three replications in RCB design having normal and late
planting in all the three situations. In 1992, thirty-day old seedlings were transplanted
in irrigated field and forty three-day old seedlings were transplanted in rainfed field
each with 5m x 2 rows plot in three replications in RCB design only at BRRI, Rajshahi
152
Fig. 2.
station. In 1993, thirty-day old seedlings were transplanted under rainfed condition
in 4m x 2 rows plot at Alimganj with three replications following RCB design. The
seeds were sown in late July to face the drought stress. Two-three seedlings were
planted per hill at a spacing of 25 x 15 cm with a fertilizer dose of 80:60:40 kg NPK/ha
for all three years. Total quantity of phosphorus and potassium were applied as
basal. Nitrogen was applied in two equal splits- one at tillering stage and another
at panicle initiation stage. Insect pests were controlled as and when necessary. Soil
moisture content from 7th October to harvesting stage was also recorded (Fig. 3).
153
Fig. 3.
Table 1
Textural Class
Sand (5%)
Silt (%)
Clay (%)
Water holding capacity (%)
Bulk density (g/cm3)
Soil pH
Organic matter (%)
Total N (%)
Available P (ppm)
Exchangeable K (meq/100g)
Available S (ppm)
Available Zn (ppm)
(01NHC1)
Cation exchange capacity
(meq / 100 g)
154
Silty Clay
5.00
51.00
44.00
57.00
1.58
5.8
1.48
0.056
15.00
0.23
15.00
2.3
15.00
Advanced lines
Highland
Lowland
Sterility percentage
Irrigated
Highland
Lowland
Irrigated
1st
2nd
1st
2nd
1st
2nd
1st
2nd
1st
2nd
1st
2nd
set
set
set
set
set
set
set
set
set
set
set
set
BR4548-2B-70
0.955 0.869
1.005
BR2575-2B-62-1-1
1.022 0.928
1.088
28.34 71.00
BR4970-37-1-2
0.991 0.882
0.935
28.48 63.52
BR4970-107-2-6
1.036 0.960
1.094
34.79 39.37
22.10 66.66
BR4974-23-1-1
1.066 0.874
1.194
BR4974-45-9-2
1.079 0.874
1.188
3211 32.56
BR4548-2B-222
0.825 0.913
1.058
16.93 38.66
BR5437-2B-13-2
1.034 0.947
1.038
13.13 42.80
BR4974-13-2-9
0.884 0.862
0.941
21.19 97.28
BR5204-B-24
0.994 0.983
0.974
17.72 46.70
BR5323-B-52
1.055 0.843
1.075
16.24 25.22
BR1870-88-1-1
0.947 1.015
0.943
16.40 28.69
BR11
0.975
0.950
0.938
44.49
24.84
26.86
BR22
0.894
0.833
0.873
48.19
54.58
22.06
Nizersail
0.959
1.066
0.974
43.62
34.68
16.47
Date of Seeding:
Date of Transplanting :
Month
Normal planting
Late planting
17.7.91
14.8.91
6.8.91
17.9.91
Temperature
Minimum (range)c
Maximum (range)c
October
19.0-25.5
25.0-35.0
November
10.0-19.0
26.0-30.5
December
7.5-13.5
17.2-28.0
155
All the twelve entries showed good panicle exertion both in the normal (1st
set) and late planting (2nd set) in case of high land and low land situation compared
to irrigated plot. The spikelet sterility of all the twelve entries of the normal planting
(1st set) under high land, lowland and irrigated situations was lower compared to
check entries. BR2575-2B-62-1-1, BR4974-23-1-1, BR5437-2B-13-2 and BR4974-13-2-9
showed exceptionally high spikelet sterility in the irrigated situation.
In 1992, twelve lines were selected on the basis of visual panicle exertion and
visual fertility score under field condition (Table 2b).
Table 2b.
Advanced
(cm)
Date of
selected
Panicle exertion
flowering
score
Fertility
score
lines/
varieties
Irrigated Rainfed
Irrigated Rainfed
Irrigated Rainfed
Irrigated Rainfed
BR4761-3B-7-2
104
62
14/11
16/11
BR4761-3B-108-2
112
61
14/11
12/11
BR4764-3B-3-2
126
72
12/11
14/11
BR4972-92-1-3-1
100
56
12/11
14/11
BR4972-92-1-3-1
100
52
12/11
14/11
BR4761-3B-208-1
100
61
12/11
14/11
BR4761-3B-18-4
106
62
13/11
14/11
BR4961-3B-70
113
85
8/11
15/11
BR1192-2B-10-1-1
86
58
BR1867-26-3-1-1
92
63
14/11
13/11
16/11
16/11
1
3
3
3
1
1
5
3
BR850-22-1-4
88
68
10/11
82
17/11
3
2
103
11/11
18/11
BR4974-48-2-3
Nizersail (ck.)
94
78
12/11
14/11
BR22 (ck.)
BR23 (ck.)
85
83
63
51
12/11
14/11
14/31
16/11
3
5
3
5
1
5
3
5
Plant height and date of flowering were also considered. All the selected
entries showed fertility score of 1, except BR4961-3B-70 and BR974-48-2-3 which
showed score of 3 and 2, respectively, under irrigated conditions. However,
fertility score under rainfed condition varied between 3 and 5 for all entries
including checks.
156
Table 2c.
List of the advanced breeding lines and varieties selected for drought
tolerance at reproductive phase. T. Aman, 1993.
Plant
Date
Panicle
height
of
exertion
(cm)
flowering
score
BR4872-72-1-4-3-1
128.2
11/11
BR4872-72-1-4-3-3
141.6
11/11
BR5226-21-2-3-2
126.6
6/11
BR5296-29-1-7-2
119.2
16/10
BR5439-39-8-1
119.7
23/10
BR5800-9-1
124.2
31/10
BR1725-13-7-1-6
105.4
21/10
BR4974-2-6-1-3-1
143.2
30/10
BR5204-213-2-9
128.4
27/10
BR5226-21-6-3-3
122.2
30/10
BR5225-2-3-1-4
120.1
20/10
BR5323-B-73-2-2
141.8
25/10
BR5452-6-2-2
123.4
10/ 10
BR5455-16-1-1
130.4
22/ 10
BR5479-11-6-1
119.1
26/10
BR554-156-1-2-1-1-519
103.1
2/11
BR5226-5-4
125.0
3/11
BR830-22-1-4
106.4
2/11
BR11(ck.)
106.3
23/10
BR22(ck.)
114.3
7/11
Batraj (ck.)
130.4
3/11
Advanced lines/
varieties
Fertility
score
In 1993, 18 lines were selected on the basis of visual panicle exertion and visual
fertility score. List of the selected entries is presented in Table 2c. Most of the entries
flowered earlier than BR22 (photoperiod sensitive modern variety). However, only
three entries BR4872-72-1-4-3-1, BR4872- 72-1-4-3-3 and BR-5226-21-2-3-1 flowered
at more or less same time with BR22. These three lines had better panicle exertion
and fertility score than BR22 and assumed to have some degree of tolerance to
drought stress at reproductive phase.
157
Fig. 4.
158
maturity and growth duration compared to the checks (BR14 and Biharibatraj),
IR54104-81-2-3-2 with an average yield of 3.83 t/ha having long slender grain was
selected which can substitute BR14 in this region.
Table 3.
Advanced
Plant
Date
Pancile
Spikelet
Yield
Phenotype
height
of
exertion
sterility
(t/ha)
acceptability
(cm)
flowering
ratio
lines
Irrig- Rainated
Irrig-
Rain- Irrig-
fed
ated
fed
at maturity
Rain- Irrig-
Rain- Irrig-
Rain-
Irrig- Rain-
ated
fed
ated
fed
ated
fed
ated
fed
BR4761-3B-7
145
138
8/11
8/11
1.02
1.01
21
38
4.06
3.35
BR4761-3B-7
145
138
8/11
8/11
1.02
1.01
21
38
4.06
3.35
BR4761-3B-108-2
112
110 30 / 10
30 /10
1.01
1.01
17
20
2.93
2.75
BR4761-3B-208-1
121
99
8/11
9/11
1.14
1.11
21
31
3.46
2.70
BR4761-3B-18-4
127
104
7/11
8/11
1.06
1.04
30
45
3.33
2.30
BR4764-3B-3-2
140
118
9/11
10/11
1.08
1.03
23
29
2.74
2.18
BR4972-92-1-3-1
124
113
6/11
5/11
1.10
1.07
24
31
3.25
2.63
BR4761-92-1-3-1
132
115
6/11
5/11
1.10
1.08
34
33
2.93
2.50
BR4761-38-70
149
124
7/11
6/11
1.06
1.05
36
25
2.27
2.12
BR1192-2B-101-1
148
122
9/11
6/11
1.03
1.01
29
37
2.88
2.43
BR1867-26-3-1-1
135
118
4/11
2/11
1.02
0.97
30
46
3.20
2.12
BR850-22-1-4
151
121
4/11
31/10
0.97
0.94
26
33
3.17
1.95
BR4974-48-2-3
131
110
7/11
4/11
1.02
0.99
22
26
3.55
3.50
BR2575-2B-62-1
117
105
7/11
4/11
1.07
1.06
24
27
3.66
2.13
BR4970-47-1-6
118
104
6/11
5/11
1.02
1.02
28
37
3.60
3.25
BR4974-23-1-1
140
115
3/11
4/11
1.06
1.03
34
28
3.55
2.95
BR4974-34-6-6
149
117
5/11
5/11
1.03
0.98
31
36
3.71
3.00
BR4974-42-1-3
154
117
4/11
4/11
1.03
1.01
27
32
4.26
3.48
BR4974-45-9-2
148
114
4/11
5/11
1.11
1.05
31
27
3.87
3.46
BR11 (ck.)
111
95 24/10
20/10
0.96
0.97
28
28
4.34
3.25
BR22 (ck.)
122
101
10/11
7/11
0.93
0.94
28
29
4.19
2.95
Niwrsail (L.ck.)
144
125
8/11
4/11
1.01
1.09
17
29
3.17
2.86
160
Table 4.
Distribution (%)
Advanced lines
0-10
10-20
20-30
0-10
10-20
20-30
BR4761-3B-7
2.364
.033
.012
98.13
1.37
0.50
BR4761-3B-20-2
2.418
.041
.014
97.66
1.65
0.69
BR4761-3B-208-1
2.447
.059
.034
96.34
2.32
1.34
BR4761-3B-18-4
3.086
.049
.019
97.85
1.55
0.60
95.74
2.96
1.30
BR4764-3B-3-2
2.810
.087
.038
BR4972-92-1-3-1
2.345
.059
.024
96.58
2.43
0.99
BR4772-92-1-3-2
2.466
.044
.019
97.51
1.74
0.75
BR4761-3B-70
2.412
.095
.022
95.37
3.76
0.87
BR1192-2B-10-1-1
2.823
.067
.028
96.74
2.30
0.96
BRl867-26-3-1-1
2.973
.077
.022
96.78
2.51
0.71
BR850-22-1-4
3.115
.059
.016
97.65
1.85
0.50
BR4974-48-2-3
2.842
.051
.027
97.33
1.75
BR2575-2B-62-1-1
2.728
.075
.038
96.02
2.64
1.34
BR4970-47-1-6
2.435
.085
.019
95.90
3.35
0.75
BR4974-23-1-1
2.356
.063
.039
95.85
2.56
1.59
BR4974-34-6-6
2.772
.059
.038
96.62
2.06
1.32
.079
.017
95.41
3.78
0.81
92.37
BR4974-42-1-3
1.994
0.92
BR4974-45-9-2
1.961
.132
.030
6.22
1.41
BRll (ck.)
2.002
.063
.020
96.02
3.02
0.96
BR22 (ck.)
3.133
.072
.012
97.39
2.24
0.37
Nizersail (ck.)
2.607
.057
.050
96.06
2.10
1.84
Due to undulating landscape, sampling was done from high and low toposequences
dividing each into 3 sites and from each site 10 sampling spots were chosen.
Table 5.
Advanced lines
Plant
height
Panicle
m/ 2
(cm)
Phenotypic
acceptability at
Days
to
Days
to
Grain
yield
Veg.
Mat.
flow.
mat.
(t/ha)
BR5000-2B-6-1
143
188
100
127
2.0
BR5344-38-2-2
121
178
112
139
2.3
BR5363-10-2-1
105
244
92
119
3.6
BR5363-10-2-3
112
256
92
119
3.7
IR28941-1-3-5-1-2
82
224
106
132
2.9
IR38499-60-368-3-21-1
95
177
95
121
2.7
IR40931-26-3-3-5
106
194
107
133
3.2
IR41054-81-2-3-2
97
260
93
119
3.7
IR48776-10-2-1
104
246
93
120
2.3
B3894-40D-PN-5-1
109
204
97
124
2.9
B5986-MR-B-1-10
99
234
94
121
3.1
102
234
96
124
3.8
BR14(ck.)
Date of sowing : 10-7-92;
Table 6 :
163
Table 7.
10-20
20-30
0-10
10-20
20-30
Low toposequence
Site 1
5.94
0.031
0.004
99.41
0.52
0.52
Site 2
6.85
0.037
0.002
99.43
0.54
0.03
Site 3
9.48
0.063
0.002
99.32
0.66
0.02
High toposequence
Site 1
6.20
0.030
0.002
99.49
0.48
0.03
Site 2
6.51
0.042
0.001
99.34
0.64
0.02
Site 3
8.70
0.046
0.003
99.44
0.53
0.03
164
Table 8.
(mg/cm 3 )
Ploughpan management
V1
V2
V3
V4
V5
4.20
3.28
3.94
3.18
(97)
(98)
(98)
(98)
(98)
4.09
4.25
3.32
3.88
3.19
deep tillage
(96)
(96)
(97)
(96)
(96)
3.38
4.16
3.32
3.95
3.03
deep tillage
(95)
(95)
(97)
(95)
(95)
4.10
4.32
3.45
4.05
3.26
(95)
(96)
(96)
(96)
(96)
0.09
0.05
0.08
0.07
(3)
(2)
(2)
(2)
(2)
0.16
0.18
0.10
0.15
0.14
(4)
(4)
(3)
(4)
(4)
0.21
0.23
0.12
0.21
0.16
(5)
(5)
(3)
(5)
(5)
0.23
0.22
0.14
0.14
0.15
(50)
(5)
(4)
(4)
(4)
**
165
Table 9.
Ploughman management
V1
V2
V3
V4
V5
Mean
2.9
3.3
3.9
4.3
4.6
4.9
4.7
5.0
5.2
5.2
5.3
3.1
3.5
4.5
5.3
5.2
5.6
2.7
3.5
4.5
5.0
0.63
5.0
5.3
2.8
3.3
5.9
6.1
5.4
5.8
6.1
5.9
6.0
6.1
5.8
6.1
0.25
5.7
4.5
3.9
5.3
5.5
5.9
4.8
5.7
5.7
5.5
5.0
5.6
5.8
5.4
4.8
Table 10.
Ploughpan Management
by
ploughpan
10
15
20
25
30
0.5
2.0
2.2
2.2
1.5
1.0
1.5
1.5
1.0
0.1
0.1
1.0
1.5
1.0
0.5
0.7
0.7
1.0
1.2
0.5
0.5
166
0.5
REFERENCES
BBS (1992) Bangladesh Bureau of Statistics, Dhaka, Bangladesh.
BRRI (Bangladesh Rice Research Institute) (1992) regional station, Rajshahi research report for 1991 wet
season, Rainfed lowland rice research consortium site at Rajshahi, Bangladesh.
Haque M E, Chang T T, Tepora N M, G C Loresto (1988) Root and shoot characters of rice ( Oryza sativa
L.) in relation to drought resistance aeroponic and hydroponic culture. Bangladesh J. Plant Breeding and
Genetics 1 (l&2): 18- 25.
IRRI (International Rice Research Institute (1992) Challenges and opportunities in less favourable
ecosystem: Rainfed lowland rice. IRRI information series no. 1, IRRI, Los Banos, Philippines.
OToole J C, Chang T T (1978) Drought and rice improvement in perspective. IRRI Research Paper Series
No. 14.:IRRI, Los Banos, Philippines.
Manalo E B (1975) Agro-climatic survey of Bangladesh. The Bangladesh Rice Research Institute and the
International Rice Research Institute.
Russell M B (1959) Water and its relation to soils and crop. Advances in Agron. 11 :1-122.
167
SUMMARY
A soil-water balance module (PADDY) and a module predicting the response
of rice to drought (DSTRESS) were developed and linked with a crop growth
simulation model (ORYZA1) to simulate rice growth and production in rainfed
lowland rice ecosystems. The combined PADDY-DSTRESS-ORYZA1 model
(ORYZA-W) was validated using two field experiments in the Philippines. Measured
and simulated changes in ponded water depth under flooded soil conditions were
in good agreement. In one of the field experiments, temporary drought was induced
at different stages. The model satisfactorily predicted change in root zone water
content, leaf area index, total aboveground dry matter, and panicle dry weight across
drought treatments over time. The model was subsequently used to predict rainfed
rice yield as a function of soil hydraulic properties and long-term weather data (25
yr) in Tarlac Province of Philippines. Risk involved in growing rainfed rice was
quantified by calculating yield probability distribution for seven major soil types
under rice cropping. Coupling ORYZA-W to a GIS allowed a spatial analysis of risk
in Tarlac Province.
INTRODUCTION
Process-based simulation models can be used as a tool to unravel some of the
complexity and variability of rainfed lowland rice ecosystems. Such models allow
detail analysis of experimental data, or extrapolation of research findings to other
environments.
Quantifying the responses of rice to drought stress is essential for predicting
the impact of soil and weather conditions on rice production. For lowland rice, grown
in puddled soils, there is little information on the link between soil-water status and
crop response, although drought is generally seen as a major cause of yield loss in
rainfed rice production system. Existing rice growth simulation models use standard
drought stress responses often derived for other crops.
The man-made puddled layer in lowland rice soils is often effective reducing
water loss through percolation to deeper soil layers. The effect of puddling on the
1 Department of Theoretical Production Ecology, Wageningen Agricultural University, P.O. Box 430, 6700 AK
Wageningen, The Netherlands
2 International Rice Research Institute, P.O. Box 933, 1099 Manila, Philippines.
hydraulic conductivity of the various layers is, however, not well understood. Drying
of previously submerged rice soils creates cracks that may extend through the plow
sole at the bottom of the puddled layer. This can cause a drastic and often irreversible
increase in water losses due to increased percolation rates. Existing soil-water balance
modules do not consider such changes and are not directly applicable to puddled
rainfed lowland rice soils.
A new soil-water balance module, PADDY, and a drought stress module,
DSTRESS, were developed and coupled to the model ORYZAI, (Kropff et al., 1993;
IRRI, 1993) for use in rainfed ecosystems. The combined ORYZAI-DSTRESS-PADDY
simulation model (ORYZA-W) was validated using two experiments conducted at
IRRI in 1991 and 1992, and used to predict rainfed rice yield on a regional scale for
a province in the Philippines.
Geographical latitude ;
Plant density ;
Parameters
describing
the
morphological
characteristics of the rice variety.
and
physiological
169
Fig. 1 :
The continuous drying of a puddled clay soil results in shrinkage cracks and
subsidence of the soil surface. To simulate cracking of the puddled root zone,
knowledge of the soils shrinkage characteristic is needed (Fig. 2), where moisture
ratio v is defined as the volume of water Vw over the volume of the solid phase Vs,
and void ratio e is defined as the volume of pores Vp over the volume of solid phase.
The shrinkage characteristic is used to calculate the volume of pores per volume of
soil (e, m3 /m3 ), the volume of water per volume of soil (q, m3 /m3 ), the subsidence
of the puddled soil surface, and the change in crack volume (Bronswijk, 1988). It is
assumed in PADDY that shrinkage is irreversible and that the puddled muddy
topsoil gradually regains structure, a process that usually is referred to as soil
ripening. Total porosity e therefore, declines upon drying and will not increase if
the water content of the root zone increases. In PADDY, cracks are assumed to have
penetrated through the plow sole if its simulated moisture ratio drops below 1.2,
which for IRRI soil is equivalent to a soil pressure potential h of - 100 kPa (IRRI,
1992).
Fig. 2 .
Soil shrinkage characteristic of a puddled clay soil. Values in diagram indicate soil pressure
heads (cm).
If soil cracks have not yet reached the plow sole, it is assumed that all incoming
water is used to replenish the first soil compartment but that cracks will not close.
During this phase the percolation rate is zero. As soon as the water content of the
top compartment has reached saturation, water starts ponding again and the
percolation rate will be governed by the hydraulic conductivity of the plow sole.
The amount of water that can be stored in the top compartment is calculated taking
into account the changes in volume and porosity of the top compartment due to
cracking. If cracks are deep enough to reach the plow sole (i.e.h<100 kPa), all water
in excess of field capacity will be drained from the top compartment. Because of the
171
soil ripening process, the conventional field capacity concept (i.e. volumetric water
content at h=-10 kPa) is hard to use for puddled soil conditions. The field capacity
water content of the topsoil was, therefore, defined as 95% of total porosity. For the
nonpuddled subsoil, the conventional definition for field capacity was used. In
PADDY, water that drains from the cracked root zone will fill up soil layers below
the root zone up to field capacity. Any excess water will be drained at a maximum
rate equal to the saturated hydraulic conductivity of the subsoil horizon. The water
content of soil compartments below the groundwater table depth is reset to
saturation.
The required data for PADDY are :
*
Description of DSTRESS
DSTRESS is largely based on data from a greenhouse experiment conducted
at IRRI in the dry season (December - May) of 1992. In this experiment, the response
of lowland rice cultivars IR20 and IR72 grown in puddled clay soil to temporary
drought at different growth stages (transplanting, 2 wk after transplanting, midtillering, panicle initiation, and flowering) was studied. Morphological responses
(inhibition of leaf production, leaf rolling, appearance of dead leaves) and
physiological responses of the crop (reduction in transpiration rate and decrease in
development rate) were expressed as a function of soil moisture ratio of the root
zone. A similar approach was taken by Sinclair (1986) and McCree and Fernandez
(1989). During the experiment, the degree of leaf rolling was visually examined daily
at midday using a leaf rolling scale (1-5). A leaf score of 1 indicates the fipst sign of
leaf rolling, whereas a score of 5 means that the leaf has completely rolled up (after
O, Toole and Cruz, 1980). The experiment was repeated in the 1992 wet season. For
reasons of brevity, only results from the drought at mid-tillering treatment conducted
in the dry season are reported here (Fig. 3).
172
Fig. 3 .
Relationships between soil moisture ratio (cm 3 /cm-3 ) and (a) relative leaf growth, (b) leaf score,
(c) % dead leaves, and (d) relative transpiration rate in a greenhouse experiment conducted
at IRRl in the dry season of 1992 (from Wopereis, 1993).
Relative leaf growth (defined as the ratio between the leaf growth of stressed
plants and that of well-watered plants) decreased rapidly from 1 to 0 if soil moisture
ratios dropped below 1.7, i.e. leaf expansion of plants subjected to drought stopped
(Fig. 3a). Leaf rolling started at lower moisture contents, and leaf rolling score
increased from 1 to 5 within a relatively narrow range of soil moisture ratios (Fig.
3b). As drought progressed, the percentage of dead leaves increased rapidly as well
(Fig. 3c). Both leaf rolling score and percentage of dead leaves were linearly related
to soil moisture ratio. Transpiration rate per unit of area of plants subjected to drought
(Td) remained equal to that of well-watered plants (Tw), even if soil-water status
dropped nearly 50%. As soil-water content declined further, a decrease in relative
transpiration rate (defined as Td/Tw) was observed (Fig. 3d). Well-watered plants
and plants that were temporarily stressed in the vegetative phase did not differ
173
significantly in yield for either cultivar. However, flowering and maturity were
strongly delayed. Severe drought in the reproductive phase greatly reduced yields.
The following morphological and physiological plant responses to drought were
quantified for the different growth stages : a) inhibition of new leaf production, b)
leaf rolling, c) leaf senescence, d) decrease in relative transpiration, and e) decrease
in development rate in the vegetative stage. Responses a, b, and c followed each
other more or less sequentially, whereas response d) started at roughly the same
soil-water content as response a) and declined to zero when leaves were dead (end
of c) (Fig.3).
The effect of drought stress on development rate in the vegetative phase (e)
could not be directly measured and is therefore not shown in Figure 3. In DSTRESS,
the development rate in the vegetative phase used in ORYZA1 (Kropff et al., 1993)
is multiplied by a factor that increases from 0 to 1 between transplanting and
flowering. This means that the closer the development stage is to flowering, the
smaller the postponement effect. No delay in growth is simulated if drought occurs
in the reproductive phase.
In the greenhouse study, production of new leaves was strongly inhibited than
dative transpiration per unit leaf area during drought periods in the vegetative phase.
This means that CO2 assimilation continues, but the C produced cannot be used for
leaf production. In DSTRESS, excess C is stored in a pool and released for leaf production
as soon as drought stress is released. The 1992 dry season experiment was repeated in
the 1992 wet season.
Model validation
The ORYZA-W model was tested for flooded soil conditions using data from
a field experiment conducted at IRRI in the 1991 dry season (cv IR72) and described
in detail by Wopereis et al. (1994) and Bouman et al. (1994). Input variables were
rainfall, irrigation, evapotranspiration rate from daily weighing of pots installed in
the field, and groundwater table depths measured using piezometers. Average and
upper and lower extreme values for measured hydraulic conductivity of the plow
sole and the non puddled subsoil were used. Simulated and observed changes in
ponded water depth were compared.
For nonflooded soil conditions, the ORYZA-W model was tested using data
from a second experiment (field experiment 2) conducted in the 1992 dry season on
a 2,000 m2 field (cv IR72). Four drought treatments in four replications were tested
in a randomized complete block design in 10 x 5 m subplots. These subplots were
separated by bunds and hydraulically isolated by plastic sheets placed 0.6 m into
the soil. Prior to the start of the experiment, all plots were submerged for 10 d, then
plowed three times and harrowed three times using a water buffalo.
Drought was initiated at transplanting (T1), at mid-tillering (T2), at panicle
initiation (T3), or during the grain filling stage (T4) by simply draining the ponded
water from the plots. For comparision, a well-watered treatment was included (T0).
Two drought durations (D) were imposed based on the 1-5 leaf rolling scale of
O'Toole and Cruz (1980). For Dl, plots were rewatered to allow plant recovery from
174
drought when leaves showed initial leaf rolling (leaf score = 1). For D2, plots were
rewatered when leaves showed clear sign of leaf rolling (leaf score = 3).
All simulations were conducted using parameters obtained from the 1992 dry
season experiment with IR72 reported in Kropff et al. (1993), except for the
development rates of the vegetative and reproductive phase and leaf N concentration
as a function of time, which were derived from the well-watered TO plots.
Model application
The ORYZA-W model was used to predict rainfed rice yield for the wet season
(June-November) in the Province of Tarlac which is located in the northern part of
the Philippines on the island of Luzon (Fig. 4). It covers an area of approximately
300, 000 ha and comprises 17 municipalities with a total population of about 740,
000 (BSWM, 1992).
A soil map of Tarlac Province (1:50, 000), which was provided by the Bureau
of Soils and Water Management, Quezon City, Philippines, was digitized using the
geographic information system (GIS). The total number of mapping units was
reduced from 67 to 14 through generalization taking into account similarity in soil
properties and importance of the unit in terms of surface area. A similar approach
was taken by Bregt et al. (1989).
As a first qualitative step, soil unsuitable for rice growth, which included the
mountainous area and light-textured soils, were eliminated from the analysis.
However, light textured soils classified on the soil map as severely flooded because
of their proximity to a river were not excluded; it was assumed that rice grown on
such soils does not suffer from drought stress. Simulations were conducted for
potentially suitable soils only. Potential (irrigated) rice yield was simulated using
the model ORYZA1 for 25 yr of weather data derived from a meteorological station
in the center of the province.
Water-limited (rainfed) rice yield for the same set of 25 yr was simulated using
the ORYZA-W model. Guided by the detailed soil map representative profiles for major
soil type under rice cropping were located. The soil hydraulic properties needed in
ORYZA-W were determined for all soil horizons in each representative profile.
Procedures and results of these measurements are presented elsewhere (Wopereis et al.,
1993). Crop parameters for rice cultivar IR72 were derived from a wet-season experiment
conducted at IRRI in 1991 (Kropff et al., 1993). Simulations started at transplanting,
assuming that seedlings were 30 days old. Initial leaf area index (LAI), temperature sum
(sum of thermal units over a base temperature), and development stage of the seedlings
were taken from field experiment 2. Initial rooting depth was assumed to be 0.05 m.
After discussion with an expert from the Bureau of Soils and Water Management,
Quezon City, Philippines (W. Sanidad, pers. commun.), transplanting of rice was
assumed to start when cumulative rainfall exceeded 75 mm during seven consecutive
days after 1 June. Thickness of the puddled topsoil was set to 0.15 m with the plow sole
occurring between 0.15 and 0.20 m depth. At transplanting, the puddled topsoil was
assumed to be saturated with an initial ponded water depth of 0.05 m. Subsoil horizons
were assumed to be at field capacity (h= -10kPa).
175
Fig. 4.
layer (i.e. plow sole) in the top 0.2 m of a puddled clay soil at the experimental farm
of the International Rice Research Institute. Average value was 0.036 cm/d, with
95% confidence limits at 0.027 and 0.045 cm/d. In this study, two classes of puddling
(poorly puddled and well puddled) were considered and expressed in terms of the
hydraulic conductivity ks of the plow sole: well puddled, Ks (plow sole) = 0.01 cm/d
and poorly puddled Ks (plow sole) = 0.10 cm/d. Combined with the two water table
depths, four simulation series were created:
1 :
2 :
3 :
4 :
Probability distributions of rainfed rice yield for all soil types under rice
cropping were estimated. Maps of simulated rainfed rice yield at different levels of
cumulative probability were produced using the GIS software.
Fig. 5.
Simulated and observed changes in ponded water depth in field experiment 1 using soil-water
balance module PADDY.
177
Fig. 6.
178
Simulated (lines) and observed (symbols) total dry matter weights (t ha-1) for cv. IR72 in field
experiment 2 for all drought treatments : A is drought at transplanting, early recovery; B at
transplanting, late recovery; C at mid-tillering, early recovery, D at mid-tiliering, late recovery;
E at panicle initiation, early recovery; F at panicle initiation, late recovery; G at flowering, early
recovery; H at flowering, late recovery.
complicated differential SAWAH soil-water balance module (ten Berge et al., 1992)
using the same field data. Results from this study showed that the iteration procedure
used in PADDY to calculate the flux through the soil profile under flooded soil conditions
was as effective as the small time step calculations used in SAWAH. The observed and
simulated total above ground matter (Fig. 5), LAI, panicle dry weight, and root zone
water content (Fig. 6) were compared. The results indicated that ORYZA-W could
satisfactorily explain the differences in biomass production and soil water content across
drought treatments, although for root zone, water content predictions were less good
at severe stress and for drought at flowering.
Tarlac simulations
Potential yields in Tarlac Province varied from 5.4 to 6.7 t/ha (Fig.7), which
are considerably higher than the irrigated rice yields (2.5-3.5 t/ha) reported by BSWM
(1992). This discrepancy may be due to a number of factors, e.g. lack of fertilizer,
incidence of pest and diseases, etc. that wen' taken into account by the ORYZA-W
model.
Fig. 7.
Simulated (lines) and observed (symbols) soil water content m 3 /m -3 for cv. IR72 in field
experiment 2 for drought at transplanting (0-5 cm, late recovery, closed circles), drought at
mid-tillering (0-10 cm, late recovery, squares), drought at panicle initiation (0-10 cm, late
recovery, triangles), and drought at flowering (0-1 0 cm, no recovery, open circles).
Rainfed rice yields ranged from 0 to 6.7 t/ha. For reasons of brevity, the
variability of rainfed rice yield over 25 years for the four simulation series is shown
in Fig. 8 for two distinct soil types only : Zaragoza clay loam (light texture) and
Padapada clay soil series (heavy texture). Potential yields are also shown for
comparison.
179
Fig. 8.
180
Potential and rainfed rice yield calculated for 25 consecutive wet seasons and four simulation
scenarios on Zaragoza and Padapada soil.
Fig. 9.
Cumulative distribution functions for rainfed rice yield on Zaragoza and Padapada soil.
181
Comparison of rainfed rice yield with potential yields quantifies the yield gap
between fully irrigated and rainfed production. This information indicates the yield
loss farmers will experience due to lack of irrigation water, under otherwise optimal
growing conditions. Production risk was quantified by calculating cumulative
probability functions for rainfed rice yield for each soil type (Fig. 9). For Zaragoza,
a shallow groundwater table had a positive effect on grain yield due to increased
capillary rise to the root zone. For Padapada, this effect was almost non- existent.
Poor puddling resulted in yield losses for both soils, especially for Zaragoza.
The hydraulic conductivity of the plow sole was an important determinant
of rainfed rice yield for light-textured soils with a relatively permeable subsoil, like
the Zaragoza soil series. If no information on this soil parameter is available, a
constant percolation rate determined for the various soil types may be used as an
input for the PADDY soil-water balance module (Wopereis et al., 1994; Bouman et
al., 1994). Tuong et al. (1994) showed that percolation losses toward and into bunds,
and the effect of poorly puddled sites may be important in areas with a relatively
permeable subsoil. More complex numerical models that allow for lateral flow into
the bunds (eg. Walker and Rushton, 1984) are needed under these circumstances.
On a regional scale, one-dimensional models, such as the PADDY soil-water balance
module can still be used, provided a constant percolation rate is assumed,
incorporating both vertical and lateral percolation losses.
Simulated rainfed rice yield was mapped at the 10 and 90% cumulative
probability levels using the GIS software. Simulations were conducted assuming
average ks values of the plow sole determined in the laboratory and a water table
depth of 1.0 m. The Zaragoza soil series occupies a large part of the potential
rice-growing area in Tarlac Province (Wopereis, 1993). Growing rice under rainfed
conditions in that province is, therefore, risky.
The approach outlined above can only result in a broad overview of yield
losses due to drought in Tarlac. Soil types were characterized using measurements
conducted at one representative site only. Spatial variability of soil hydraulic
properties or thickness of soil horizons was not taken into account.
CONCLUSION
The GIS and crop simulation modelling can be used to quantify rice yield
losses due to drought at a regional level. Rainfall variability has a strong impact on
yield variability in Tarlac Province. Field experiments conducted for 1 or 2 yr in such
environments may give misleding results. Long-term weather data are needed to
determine probability distributions of crop yield to perform an economic evaluation
(Anderson, 1991). Unfortunately, there is a lack of long-term weather data in many
rice-growing countries in Asia as was also the case for the study presented here
(detailed weather data were only available for one station in the entire province).
Supplementary irrigation increased wet-season rainfed rice yields and
reduced yield variability. Irrigation may also increase the potential for a dry-season
crop (eg. mungbean), which would boost total production and income per year
relative to rainfed conditions. The scope for a dry-season crop after rice could be
182
investigated using the PADDY soil-water balance and good explanatory model for
the dry-season crop.
FUTURE RESEARCH NEEDS
One of the major problems in the application of crop-soil models is the lack
of data on soil hydraulic functions. Data bases, that relate these functions to soil
characteristics derived from soil survey data, are needed. Maintenance and
installation of weather stations should be promoted partly through research
consortia. It is important to have a few well-selected and well-maintained sites that
can be carefully monitored. Only if good data sets are available can simulation
models be used to extrapolate new technologies or to identify constraints or
opportunities in rice production. More research is needed on how to deal with limited
data in crop modelling studies.
The rainfed rice model ORYZA-W can be used to investigate yield losses due
to drought in rainfed, puddled environments and may also be used to quantify the
benefits of improved irrigation facilities. ORYZA-W is also a starting point for
simulation studies on rice upland crop rotations. The model, however, needs further
validation for a broader range of environmental conditions and for different rice
varieties. PADDY needs to be expanded to include simulation of water table depth.
Sensitivity analyses to investigate the importance of variability in model-input
parameters must be conducted.
The increased water-use efficiency of rice-based cropping systems is becoming
increasingly important. This can be done by improving irrigation facilities,
introducing water-saving techniques, and adjusting the planting time and/or
cropping system to maximize rainfall utilization. For any of these approaches, a
thorough systems analysis to evaluate the different solutions for different
environments is needed. One of the most promising water-saving techniques is dry
seeding of rice. The drought stress functions used in this study were all derived for
puddled soil conditions. More research is therefore needed on drought stress
responses of dry seeded rice and their relation to root zone water content. Special
attention should be paid to root development and the effect of drought on root
growth.
REFERENCES
Anderson J R (1991) A framework for examining the impacts of climatic variability. Pages 3-17, In : Climatic
risk in crop production: Models and management for the semiarid tropics and subtropics. R.C. Muchow
and J.A. Bellamy, eds. CAR International, Wallinglord, UK.
ten Berge H F M, Jansen D M, Rappoldt K, Stol W (1992) The soil water balance module SAWAH : Users
guide and outline. CABO-TPE Simulation Report Series, no. 22 CABO Wageningen, The Netherlands, 78
pp. + appendices.
183
184
Wolfram S (1991) Mathematica. A system for doing mathematics by computer Second edition.
Addison-Wesley Publishing Company Inc., Redwood City, California, 961 pp.
Wopereis M C S, Wosten J H M, Bouma J, Woodhead T (1992) Hydraulic resistenace in puddled rice soils
measurement and effects on water movements Soil & Tillage Research 24: 199-209.
Wopereis M C S (1993) Quantifying the impact of soil and climate variability on rainfed rice production.
Ph.D. Thesis, Wageningen Agricultural University, The Netherlands. ISBN 90- 5485-147-3.
Wopereis M C S, Kropff M J, Hunt E D, Sanidad W, Rouma J (1993) Case study on regional application
of crop growth simulation models: Tarlac Province, Philippines. Pages 27-46. In : Bouman, B.A.M., van
Laar, H.H. and Zhaoqian, W. (Eds). Agro- ecology of rice-based cropping systems. ISBN 90- 73384-18-4.
Wopereis M C S, Bouman B A M, Kropff M J, ten Berge H F M, Maligaya A R (1994) Understanding the
water use efficiency of flooded rice field,s I. Validation of the soil-water balance model SAWAH.
Agricultural Water Management (in press).
185
SUMMARY
Dry seeded rice (DSR) provides an option to increase crop production and
intensity by utilizing early-season rainfall, escaping late-season drought and
reducing turnaround time for a post-rice crop. Experiments conducted in 1992 and
1993 at the Tarlac Rainfed Lowland Consortium Key-site evaluated and quantified
the response of DSR to drought in a rainfed lowland environment. The treatments
were different dates of seeding and varying water regimes, each having three levels.
The seeding dates were May 18, June 11, and July 8 in 1992, and May 27, June 11,
and August 4 in 1993. The three water regimes were (i) totally rainfed conditions,
(ii) initially irrigated conditions to ensure complete emergence and then rainfed, and
(iii) fully irrigated conditions. The ORYZA1 model was parameterized for DSR
through vegetative and reproductive development rates and tested to simulate DSR
growth under fully irrigated conditions. Transpiration and evaporation were
determined by daily weighing of 20x20x40cm lysimeters installed in the fields. The
final yield was not significantly affected by the various water regimes except for the
first seeding date. Under rainfed conditions, the second seeding date resulted in the
highest yield. Rice plants transpired at a potential rate as soil water content of the
mot zone decreased from saturation to field capacity (-0.03 MPa matric potential).
Below field capacity, the relative transpiration rate decreased linearly with decreasing
soil moisture. This linear decrease appeared to be independent of the fertility level
and climatic conditions. ORYZA1 satisfactorily estimated dry matter production and
yield under fully irrigated conditions.
INTRODUCTION
About 27% of the total rice area in the world has been classified as rainfed
lowland, which contributes less than 18% to global rice production. More than 95%
of the world's total rainfed lowland area is in Asia (IRRT, 1993). Even a small increase
in the productivity of these regions would add significantly to global rice production.
With the availability of modern, short-duration, high yielding varieties, dry seeded
rice (DSR) provides an option to increase the productivity in rainfed lowland
ecosystems. DSR can be sown even before the onset of the wet season, which permits
utilization of early-season rainfall (Tuong et al., 1993). Seeds of DSR remain viable
in the soil for as long as 40 days without detrimental effect on germination (Furoc
et al., 1978). The early harvest of DSR enables the crop to escape and/or minimize
the adverse effects of drought at the end of the wet season, and leaves enough time,
as well as available soil moisture and late-season rainfall, for a second crop (Saleh
et al., 1993). Since DSR is done in nonpuddled conditions, it saves on turnaround
time for post-rice crops.
The uncertainty of rainfall may expose the crop to drought of varying intensity
and duration during the growing season. This is one of the key factors limiting yields
of rainfed rice. Advancing the seeding date in DSR aggravates this problem because
the crop is established before the steady monsoon season rain. DSR, because of its
very nature of cultivation, is associated with many problems starting with crop
establishment, high weed competition and periodic dry spells. Since the soil is not
puddled, DSR represents a totally different environment for rice root development
as well as for water and nutrient uptake compared with transplanted rice (TPR).
There is a dearth of precise and systematic information on the soil-water-plantatmospheric interactions in relation to DSR. It is, therefore, essential to quantify the
effects of drought on soil-water extraction and plant performance in DSR grown
under rainfed lowland ecosystems. It will enable development and/or modification
of a model linking environmental conditions to dynamics of soil water and plant
growth which could be utilized for identifying conditions and locations suited for
DSR, the risks associated with it and the year to year yield variations.
The main objectives of this paper are :
1)
to quantify the effect of drought on the transpiration rate of dry seeded rice; and
2)
to test the ORYZA1 model, a validated crop growth model for potential
production of TPR (Kropff et al., 1993), for DSR under fully irrigated
conditions, as a first step toward a full scale modelling of DSR under rainfed
conditions.
Fig. 1.
frequency of rainfall are highly variable and uncertain, the seeding dates were varied
during the same growing season in order to expose the crop to drought at various
stages under field conditions. The treatments were three seeding dates and three
water regimes. The dates of seeding (Dl, D2, and D3) were May 18, June 11, and
July 8 in 1992 and May 27, June 11 and August 4 respectively in 1993. Water regimes
were (i) totally rainfed (IO), (ii) irrigated initially to ensure complete emergence and
then rainfed (11), and (iii) fully irrigated control (I2). All plots were separated from
surrounding plots by plastic sheets embedded 60 cm deep around each plot. The
experiment was laid out in a split plot design with water regimes as the main plot
and seeding date as subplot treatment, replicated five times. The subplot was 7.5x11.5
m. Nitrogen (120 kg/ha) was applied in four toydressed equal splits, i.e. 15 days
after emergence (DAE), maximum tillering, panicle initiation, and heading. Twenty
five kg P/ha and 45 kg K/ha were also applied. Seeds were sown at a rate of 100 kg/ha
in rows 20 cm apart and 2 cm deep. IR72 was used in both experiments. The experimental
plots were kept free of weeds and pests, following recommended practices.
Emergence count of the crop under IO was monitored daily in four 1-m-long
rows in each subplot from seeding until a constant number, i.e. final emergence, was
attained. Plant samplings from two 0.5m-long-rows in each subplot were carried out
for biomass production, leaf area index (LAI) and leaf N content. Samplings were
188
where LAIs is the leaf area index of stressed plant and LAIns is the leaf area
index of fully irrigated plants.
Since destructive sampling could not be carried out from the lysimeters until
maturity, plants were sampled from an equivalent area within the plots of fully
189
irrigated treatments to obtain LAIns at the start and at termination of stress. LAIns
and LAIs were considered equal at the initiation of stress. After the stress was
relieved, the stressed plants began transpiring at a potential rate (Tp) that was less
than the transpiration rate of well-watered plants (Tpns). The difference between Tp
and Tpns was due to the difference in the LAI of stressed and non-stressed plants
at the end of the stress period. LAIs at the end of the stress period was derived from
LAIns Tp; and Tp ns using the relation.
where ln is natural logarithm.
Daily values of LAIns and LAIs were worked out from the LAIs values obtained
at the initiation and end of stress, assuming that the change in LAI during that period
was a linear function of time. Daily CF values were then derived using equation 2.
Modelling of DSR
The ORYZAI crop growth model was tested for simulation of growth and
yield of fully irrigated DSR. ORYZAI was originally developed and validated to
simulate growth and yield of TPR rice under potential production conditions (Kropff
et al., 1993). The model requires radiation, temperature, planting dates, crop
development, and relative growth rates as well as time course of leaf N content as
input. Measured LAI can also be input; otherwise, it is predicted by the model.
Development rate parameters and leaf N content are important for simulation
of rice growth for a given environment (Kropff et al., 1993). However, the
development rate of DSR will differ from that of TPR. As a first step, therefore, the
development rate parameters of DSR were derived from the first sowing date of I2
in the 1993 WS experiment described earlier, based on seeding, flowering, and
physiological maturity dates using the program DRATES (Kropff et al., 1993).
Measured leaf N content was used as input. Simulations predicted the total biomass
and panicle dry weight of I2 treatments for all sowing dates in 1992 and 1993. LAI
was also simulated, but the results are not presented in this paper.
Fig. 2.
Cumulative rainfall during 1992 and 1993 in dry seeded rice (DRS) as a function of days after
sowing.
below the water held by the soil at wilting point (-1.50 MPa matric potential). In this
treatment, emergence was delayed by almost a week and the emergence percentage
in 1993 was 64% (Fig. 3). The emergence in 10 of the first seeding of the 1993 WS
was triggered by 69 mm of rain (8 days after seeding) and the emergence count
stabilized at a cumulative rainfall of less than 100 mm. Germination in the subsequent
seeding dates was 80% or more because of higher initial soil-water content and
greater amount of rainfall.
Table 1.
Soil water content in 0-10 cm layer at sowing time in 1992 and 1993.
Sowing date
1992
1993
18 May
11 June
8 July
0.11
0.16
0.20
27 May
11 June
4 Aug.
0.10
0.21
0.23
191
Fig. 3.
Effect of sowing date on emergence of rainfed (I0) DSR IR72. Numbers in parentheses are
cumulative rainfall totals (mm) at which emergence count became constant.
Grain yield
The final yield was not significantly affected by the various water regimes
except on the first date of seeding (Table 2). Under rainfed conditions the second
date of seeding gave maximum yields because of better water availability throughout
the growing season. Heavy rainfall, coinciding with flowering of the crop slightly
reduced yield in I2 for the second sowing date in 1992 compared with other
treatments. Tuong et al. (1993) reported similar observations of yields remaining
unaffected even though drought spells delayed emergence and other plant
phenological stages under rainfed conditions.
Transpiration ratio
For Ta/Tp, the results of the experiments conducted during 1993 DS and WS
were pooled and are shown in Figure 4 as a function of soil water content. Ta/Tp
during the DS were for 60 and 120 kg N/ha, whereas that for WS was at 120 kg
N/ha. In well-watered plants during DS, transpiration rates were 10 mm/day and
reduced to 2 mm/day in the stressed plants at the time of drought termination (at
leaf rolling score of 5). During WS, transpiration rate of well- watered plants was
192
5-6 mm/day and was reduced to 1.5 mm/day or less at the later stages of stress.
For all seasons and N levels, the rice plant was able to transpire at a potential rate
until the soil moisture was around field capacity, i.e. at a soil pressure potential of
-0.03 MPa. Beyond this soil water content, Ta/Tp decreased linearly with decrease
in soil water content, (R2 =0.9254). In a study of transplanted rice under non
nitrogen-limiting conditions, Maligaya et al. (1993) and Wopereis (1993) obtained
similar trends in the relative transpiration rate as a functiol of soil moisture ratio,
which could be translated into soil water content. In their study, Ta/Tp began
declining at a soil moisture ratio of 1.7, representing soil water content below field
capacity when water stress was initiated at mid- tillering stage for both IR20 and
IR72. The critical soil moisture at which Ta/Tp starts to decline needs to be
investigated further as it may be both crop and stage-specific as well as being
influenced by the crop establishment method.
Table 2.
Effect of seeding date and water regimes on the yield of dry seeded rice
IR72.
Seeding date*
Dl
D2
D3
*
**
Water regime
1993
I0
2.9b
3.7ab
I1
2.5b
3.4b
I2
4.0a
4.la
I0
3.8a
4.3a
I1
4.0a
3.8b
I2
3.3b
3.9ab
I0
3.8a
3.8a
I1
3.8a
3.7a
I2
4.0a
3.8a
Dl refers to 18 May in 1992 and 27 May in 1993; D2 refers to 11 June in both 1992 and 1993; D3
refers to 8 July in 1992 and 4 Aug. in 1993.
Means having a common letter in the same column are not significantly different at the 5% level by
DMRT.
Plants respond to the imposition of stress through stomatal closure and leaf
rolling. Stomatal closure reduces water loss through transpiration whereas leaf
rolling directly reduces the leaf area exposed to the environment, which also
contributes to reduced transpiration losses. Reduction in transpiration rate directly
affects the rate of photosynthesis. This relationship (Fig. 4) representing reduction
in relative transpiration rate as a function of soil water content can be used to link
a potential production simulation model to production under water-limiting
193
Fig. 4 .
Relative transpiration rate (Ta/Tp) of DSR as a function of soil water content (WS=wet season;
DS=dry season; N60=60kg Nha; N120=120 kg N/ ha).
environments through a water balance model. This could lead to a model that can
be used to quantify the effects of water stress on crop growth and yield of DSR.
Fig. 5 .
Observed (0) and simulated () values of dry matter production for all sowing dates of I2.
195
Fig. 6 .
Comparison of observed and simulated dry matter production and panicle dry weight of
well-watered DSR IR72.
tables occur frequently and for several weeks an unsaturated layer can be observed
below a saturated one in the soil profile. Initial testing of process-based water balance
model SAWAH has indicated that it can be used successfully under these situations
(Tuong et al., 1993).
196
CONCLUSION
Results of our experiments indicate that Ta/Tp was a sound parameter for
characterizing stress as it appeared to be independent of N application rate and the
prevalent climatic conditions. ORYZA1 satisfactorily simulated the dry matter and
yield of DSR under fully irrigated conditions. Further work is required to modify
the model to simulate crop growth under water- stressed situations, and coupling
the crop component with a water balance model for rainfed lowland ecosystems.
REFERENCES
Furoc R E, Magbanua R D, Gines H C, Morris R A (1978) Identification of criteria for late dry-seeded rice
planting. Paper presented at the 9th Annual Scientific Meeting of the Crop Science Society of the
Philippines, 11-13 May, 1978, Iloilo City, Philippines.
IRRI (1993) International Rice Research Institute. IRRI rice almanac, P.O. Box. 933, Manila, Philippines.
Kropff M J, Van Laar H H, ten Berge H F M (1993) ORYZA1-A basic model tor irrigated lowland rice
production : Simulation and Systems Analysis for Rice Production (SARP). International Rice Research
Institute, P.O. Box 933, Manila, Philippines.
Maligaya A R, Wopereis M C S, Kropff M J (1993) Drought stress responses of two lowland rice cultivars.
Paper presented at the 9th Federation of the Crop Science Society of the Philippines Scientific Meeting, 10-14
May, 1993, Aklan, Philippines.
O'Toole J C, Cruz R T (1980) Response of leaf water potential, stomatal resistance and leaf rolling to water
stress. Plant Physiology 65: 428-432
Saleh A F M, Lantican M A, Bhuiyan S I, Agua M M (1993) Increasing productivity of droughtprone rainfed
lowlands in Northern Luzon, Philippines. Paper presented at the Symposium of the rainfed Lowland Rice
Research Consortium, 8-13 Feb. 1993, Semarang, Indonesia.
Tuong T P, Ingram KT, Siopongco J D, Confessor R B, Boling A A Singh U, Wopereis M C S (1993) Performance
of dry seeded rainfed lowland rice in response to agrohydrology and N-fertilizer management. Paper
presented at the Symposium of the Rainfed Lowland Rice Research Consortium, 8-13 Feb. 1993 Semarang,
Indonesia.
Wopereis M C S (1993) Quantifying the impact of soil and climate variability on rainfed rice production. Ph
D thesis, Agricultural University, Wageningen, The Netherlands.
197
SUMMARY
Field experiments to quantify root growth of rice cultivars under rainfed
conditions were conducted at the Ubon Rice Research Centre, Northeast Thailand,
during the 1991 and 1992 wet seasons. Root length, root mass, and root:shoot ratio
were measured at panicle initiation and anthesis. Genotype, soil hydrology, and soil
type significantly affected root length density (RLD) and root mass density (RMD).
The RLD and RMD of genotypes at plots subjected to alternate wetting and drying
were generally higher than those at plots that were continuously flooded. Greater
RLDs and RMDs were observed in loamy sand soil than in clay soil. Root:shoot ratio
was higher in loamy sand than in clay soil irrespective of hydrology. Roots in clay
soil exhibited less root branching than roots in loamy sand. Loamy sand soil subjected
to alternate wetting and drying may be an appropriate growing condition for
evaluating rice root growth.
INTRODUCTION
Optimum root growth is required for stabilizing shoot growth in plants
subjected to drought stress. A deep and dense root system has been shown to be a
heritable characteristic, and has been assumed to be essential for allowing plants to
avoid or tolerate drought. Thus, rice breeders have attempted to incorporate
favourable root characteristics into rice genotypes for drought- prone, rainfed
lowland areas.
A measurable variation in root system characteristics of rice genotypes has
long been recognized (Yoshida and Hasegawa, 1983; O'Toole and Bland, 1988). It
has also been shown that soil physics, hydrology, and agricultural practices influence
rice root growth (Maurya and Ghildyal, 1975; Sharma et al., 1987; Thangaraj et al.,
1990; Ogunremi 1991). Rice roots, therefore, exhibit a large degree of phenotypic
plasticity. Coupled with genotype x environment interactions, root phenotypic
plasticity makes evaluation of root system characteristics difficult. The soundness
of information on rice root traits depends on whether plants were grown in an
environment where full expression of their growth potential is realized. Such
growing environments need to be identified.
1 Ubon Rice Research Centre, P.O. Box 65, Ubon 34000, Thailand.
2 Peanut CRSP Management Office, University of Georgia, Griffin, Georgia, 30233-1797, U.S.A.
3 Department of Soil Science, H.P. Krishi Vishwavidyalaya. Palampur-176062, H.P.. India
High toposequence,
nonflooded;
loamy sand,
alternate
wetting and
drying,
mostly
2)
Low toposequencc, loamy sand, flooded most of the time during the growing
season; and
3)
Six rice cultivars IR-20, KDML 105, NSG 19, RD9, Chieng Saen, and IR46 were
used in the 1992 experiment. Seedlings (30 d old) were transplanted in puddled
loamy sand soil, at high and low toposequence position. Experiments in both years
were laid out in randomized complete block design with five replications. Fertilizer
application, in both years, consisted of 25-25-25 kg/ha of N, P, and K incorporated
before transplanting and 50 kg/ha of N topdressed at panicle initiation (PI).
The genotypes used in the experiment were chosen to represent contrasts in
stature, photoperiod sensitivity, and drought susceptibility. KDML 105, (photoperiod
sensitive) and NSG19 (photoperiod insensitive) are traditionally selected, tall
cultivars, resistant to drought stress. IR20 (short stature high yielding) and Chieng
Saen (tall traditional cultivar) are known to be susceptible to drought stress. Cultivars
IR46 and RD9 are photoperiod insensitive, medium-statured, improved, and high
yielding.
Genotypes were sampled for roots at the same physiological age at PI and
anthesis. Root and shoot sampling was done on median plants at randomly selected
locations within plots. Roots were collected from four hills in each replication using
a 20 x 20 x 50 cm metal monolith sampler. Root sampling was centered over a hill.
Each soil core was sectioned into five 10 cm layers. Roots were washed free of soil
over a metal screen (1 mm mesh size). Root length was measured with a Comair
root length scanner (Commonwealth Aircraft Corporation Limited, Australia). Roots
were oven-dried at 70 for 48 h to determine mass. Root length density (RLD) and
mot mass density (RMD) were calculated as the quotient of mot length and oven
dry weight, and the volume of the soil sample. Above ground biomass was
determined in a manner similar to root mass. Root : shoot ratios (root oven dry
weight/shoot oven dry weight) were calculated from the data.
In the 1991 experiment, root form and proliferation were observed using the
pinboard technique. Pins were driven through the sampler in one hill in each
199
replication at anthesis. The monolith was carefully dug out, and removed from the
ground. Roots in the sampler were washed free of soil, allowing observation of
relative root distribution in the soil profile.
Fig. 1.
200
Root length density at anthesis stage of 3 rice cultivars at 3 toposequence positions (Pantuwan
et al 1992, unpubl).
Fig. 2.
Root length density of 6 rice cultivars (a) KDML 105, (b) MSG 19, (c) RD 9, (d) Cheing Saen,
(e) IR 20 and (f) IR 46 growth at high and low toposequence positions (Pantuwan et al 1993,
unpubl. data).
Fig. 3.
Root systems of three rice cultivars grown in (a) clay loamy sand (b) low toposequence and
(c) loamy sand high tosposequence position.
201
Root mass density (RMD) exhibited the same trend as RLD. Roots in loamy
sand produced greater RMD than the roots in clay soil (Fig. 4). Root mass density
was higher at high toposequence than at low toposequence positions (Fig. 5). Higher
RMD in the high toposequence may be related to the higher RLD observed at this
toposequence position.
Fig. 4.
Effect of soil texture on root mass density of 3 rice cultivars sown at low toposequence (Adapted
from Sharma, 1994).
Fig. 5.
Root mass density of 6 rice cultivars (a) KDML 105, (b) MSG 19, (c) RD 9. (d) Chieng Saen,
(e) IR 20 and (f) IR 46 grown at high and low toposequence positions (Pantuwan et al, 1993
unpubl. data)
202
More than 80% root length and 90% root mass were obtained in the top 20
cm soil layer, irrespective of soil hydrology (Tables 1, 2). Both RLd and RMD
decreased exponentially with the soil depths in all genotypes. This trend was similar
to previousstudies on rice roots (Pantuwan et al., 1992; Sharma et al., 1987). Genotypes
on the low toposequence position had higher percentage of roots in the top l0cm
soil layer than genotypes on the high toposequence position. This relationship was
reversed in the 10-50 cm soil layer.
Table 7.
Percent root length distribution of six rice cultivars grown under two
toposequence positions at Ubon Rice Research Centre, Thailand.
(Pantuwan et al., 1993, unpubl. data).
Root length distribution(%) at soil depth
Cultivar
0.10
(cm)
10-20
(cm)
20-30
(cm)
30-40
(cm)
0-10
(cm)
10-20
(cm)
Panicle initiation
High toposequence
KDML 105
NSG19
RD9
Chieng Saen
IR20
IR46
Mean
Low toposequence
KDML105
NSG19
RD9
Chieng Saen
IR20
IR46
Mean
20-30
(cm)
30-40
(cm)
40-50
(cm)
Anthesis
71.43
2.01
60.61
2.60
65.09
3.08
70.01
1.63
63.50
1.78
72.60
2.24
67.2
1.97
17.11
1.62
20.40
1.60
18.50
3.06
16.55
1.48
19.92
0.73
13.88
1.99
17.7
0.99
8.97
0.73
16.09
1.52
11.72
1.36
11.91
1.06
13.58
0.86
10.33
1.98
12.1
1.02
2.483
0.251
2.894
0.364
4.691
1.306
1.533
0.365
3.024
0.670
3.195
1.336
2.970
0.421
61.63
5.01
58.69
3.45
62.63
1.75
67.71
3.88
63.50
3.81
65.00
3.35
63.2
1.25
27.48
5.17
28.00
2.65
25.06
2.15
21.55
3.96
28.82
3.13
19.47
2.30
24.6
1.38
9.46
0.69
12.15
0.88
10.97
1.09
9.69
0.95
9.62
11.23
13.80
2.29
10.95
0.71
1.410
0.318
0.983
0.275
1.233
0.260
0.892
0.225
1.013
0.201
1.636
0.484
1.195
0.117
0.017
0.008
0.176
0.042
0.100
0.026
0.152
0.049
0.050
0.011
0.094
0.044
0.098
0.024
71.58
5.26
58.79
1.89
66.01
2.34
67.79
3.09
66.96
1.94
71.70
1.51
67.1
0.75
18.21
3.31
28.27
1.65
25.70
1.99
25.71
2.22
26.67
2.02
19.04
2.74
23.9
0.48
7.73
1.42
11.57
1.11
7.46
1.36
5.59
1.36
5.85
1.58
8.42
1.66
7.8
0.24
2.475
1.010
1.376
0.689
0.827
0.372
0.896
0.297
0.522
0.166
0.836
0.268
1.155
0.10
77.34
3.11
76.85
2.57
79.96
2.12
70.17
1.88
76.20
2.97
79.84
2.51
76 76
0.91
17.28
2.38
16.99
3.76
15.43
2.22
21.67
1.99
17.91
2.28
16.69
1.91
17.66
0.84
3.98
0.88
3.87
1.00
2.66
0.45
5.82
0.88
4.46
0.94
2.13
0.59
3.82
0.43
0.879
0.208
1.854
1.185
1.185
0.195
1.856
0.173
1.107
0.168
0.750
0.156
1.284
0.14
0.323
0.035
0.430
0.091
0.685
0.191
0.477
0.061
0.318
0.049
0.586
0.069
0.470
0.027
203
Table 2.
Percent mot mass distribution of six rice cultivars grown under two
toposequence positions at Ubon Rice Research Centre, Thailand.
(Pantuwan et al., 1993, unpubl. data).
Root mass distribution(%) at soil depth
Cultivar
0-10
(cm)
10-20
(cm)
20-30
(cm)
30-40
(cm)
0-10
(cm)
10-20
(cm)
Panicle initiation
High toposequence
KDML 105
NSG19
RD9
Chieng Saen
IR20
IR46
Mean
Low toposequence
KDML105
NSG19
RD9
Chieng Saen
IR20
IR46
Mean
20-30
(cm)
30-40
(cm)
40-50
(cm)
Anthesis
75.66
2.53
75.75
3.21
75.24
1.77
83.63
0.90
76.91
1.84
78.36
1.08
77.6
1.29
15.37
21.65
13.52
21.48
15.09
2.27
11.06
0.53
14.83
0.89
11.27
1.56
13.5
0.79
8.03
20.96
10.36
1.85
8.73
1.58
5.08
0.67
7.74
1.13
9.90
1.58
8.3
0.77
0.941
0.302
0.371
0.093
0.935
0.358
0.233
0.042
0.525
0.099
0.480
0.055
0.581
0.l20
77.07
0.48
73.96
2.24
73.27
1.32
75.63
2.56
78.09
1.23
76.86
2.59
75.8
0.77
17.09
1.06
18.94
1.89
19.57
0.89
18.79
2.29
17.24
0.71
16.66
2.23
18.0
0.49
5.31
0.57
6.26
0.49
6.29
1.03
5.08
0.60
3.66
5.68
1.85
1.09
5.41
0.40
0.519
0.008
0.15 0.001
0.804
0.037
0.253 0.003
0.857
0.013
0.230 0.003
0.478
0.039
0.055 0.002
0.983
0.030
0.140 0.004
0.618
0.009
0.104 0.002
0.710
0.023
0.082 0.006
83.22
2.73
72.25
1.76
75.52
2.27
80.20
1.79
76.39
0.94
82.03
1.31
78.3
1.73
12.84
1.59
20.64
1.56
20.07
1.35
16.77
1.49
19.90
1.6
14.46
1.62
17.4
1.35
3.18
1.18
6.71
0.69
4.22
1.17
2.79
0.78
3.45
0.85
3.27
0.50
3.9
0.59
0.860
0.334
0.391
0.175
0.181
0.048
0.239
0.084
0.258
0.104
0.235
0.069
0.361
0.104
83.72
2.69
78.63
1.53
82.89
1.74
85.25
1.06
85.49
1.86
82.98
2.54
82.8
0.93
14.13
1.90
16.49
2.03
14.20
2.07
13.66
0.68
12.15
1.06
14.99
2.20
14.3
0.59
1.74
0.70
3.20
0.80
2.24
0.39
2.54
10.35
1.99
0.86
1.73
0.53
2.2
0.23
0.377
0.029
0.251 0.008
0.047
1.637
0.458 0.008
0.580
0.090
0.190 0.013
0.047
0.504
0.101 0.011
0.297
0.074
0.011
0.193
0.040
0.258
0.054 0.006
0.054
0.609
0.211 0.009
root:shoot ratio (Table 3). Root : shoot ratios were higher in loamy sand than in clay
soil. High nutrient content associated with a clay texture, coupled with assured water
supply, apparently led to the production of more shoots than roots. This may indicate
that under nonlimiting nutrient and water supply, deep or extensive root systems
may not be required. Root : shoot ratio did not vary as much among genotypes as
among growing environments (see SEM values in Table 3).
Table 3.
Cultivar1
Physiological
age
High/
loamy
sand
Low/
loamy
sand
Low/
clay
type
High/
loamy
sand
1991
Low/
loamy
sand
MeanSEM
1992
Panicle initiation
KDML105
0.37
0.34
0.26
0.16
0.19
RD9
0.36
0.28
0.25
0.18
0.17
0.250.039
IR46
0.41
0.36
0.34
0.17
0.15
0.290.059
Mean
0.38
0.33
0.28
0.17
0.17
0.015
0.024
0.006
0.012
KDML105
0.14
0.13
0.09
0.09
0.10
0.110.012
RD9
0.15
0.14
0.11
0.12
0.13
0.130.008
IR46
0.14
0.13
0.08
0.09
0.12
Mean
0.14
0.13
0.09
0.10
0.12
SEM
0.003
0.003
0.008
0.010
0.009
SEM
0.0028
0.260.046
Anthesis
0.130.016
CONCLUSION
Under rainfed conditions, rice root systems were significantly affected by
genotype, soil water regime, and soil type. Differences in root systems among
genotypes were large in loamy sand soil subjected to alternate wetting and drying.
Differences in root systems decreased under flooded conditions on fine textured
soils. Root:shoot ratios were affected more by soil type than by soil hydrology.
Coarse-textured soils subjected to alternate wetting and drying may provide
appropriate growing environment for the evaluation of root systems of rainfed rice.
205
REFERENCES
Kawata S, Ishihara S (1959) Study on the root hairs in rice plant. Proc. Crop Sci. Soc. Jpn. 27: 341-348.
Kar S, Varade S B, Ghildyal B P (1979) Pore size distribution and root growth relations of rice in artificially
synthesized soils. Soil Sci. 128:364-368.
Maurya P R, Ghildyal B P (1975) Root distribution pattern of rice varieties evaluated under upland and
flooded soil conditions. (Italian summary) Riso 24: 239-244.
Ogunremi LT (1991) lnfluence of bulk density and moisture regime of a permeablesoil on the performance
of lowland rice, Oryza sativa L. Trop. Agric. 68: 129-134.
O'Toole J C, Bland W L (1988) Genotypic variation in crop plant root systems. Adv. Agron. 41: 91-145.
Sharma P K, De Datta S K, Redulla C A (1987) Root growth and yield response of rainfed lowland rice
to planting method. Exp. Agric. 23: 305-313.
Thangaraj M J, O'Toole J C, De Datta S K (1990) Root response to water stress in rainfed lowland rice.
Exp. Agric. 26: 287-296.
Yoshida S, Hasegawa S (1982) The rice root system : its development and function. Page 97-114. In :
Drought resistance in crops with emphasis on rice. International Rice Research Institute, Philippines.
206
SUMMARY
Socio-economic studies were conducted in Chandpur and Mungishpur
villages of eastern U.P., interfaced with the biophysical characterization of selected
rainfed environments. This paper examines the labour contributions by gender
differentiation and social and economic groups and the problems of women farmers
in agricultural productions in different lowland rice ecosystems. The paper provides
suggestions which can improve crop and animal productivity as well as improve
the welfare of the entire family, particularly the females, and children. The study
revealed that the sexual division of labour is related to the social position of
the household and access to non-farm income. The crop-production, livestock
raising and non-farm activities are interrelated. The farmers can bear more risks
in rice cultivation if they have access to income from non-farm sources and would
be receptive to input-intensive technologies.
INTRODUCTION
In recent years, greater attention is being given to the complex, diverse
and risk prone (CDR) rainfed regions where mostly the poor people are located.
India has 40 million hectares of rice growing areas. While Northern and Southern
India has benefitted from the modern rice varieties and irrigation, the vast rainfed
areas in eastern India has lagged behind. Eastern India which is comprised of
Assam, Bihar, West Bengal, Orissa and eastern parts of Uttar Pradesh and Madhya
Pradesh has the highest incidence of rural poverty in the country. Although it
is the largest rice growing area in India, accounting 60% of the country's rice
area (26.8 million ha), only a third of the total rice production comes from this
region. About half (480 million) of the country's population living in this region
is largely dependent on rice farming. Macro level analysis of rice growing ecosystems
showed that out of the 26.8 m ha rice areas, only 21.2% is irrigated, 47.7% is
under rainfed lowlands and the remaining 14% is under deep and very deep
water ecosystems. The rainfed lowlands ape drought prone and submergence
prone (Singh, 1992).
METHODOLOGY
We selected two typical lowland rainfed sites, in eastern Uttar Pradesh,
namely Chandpur and Mungishpur which are different in terms of proximity
to the city and physical endowments. Chandpur is closer (3 kms) to Faizabad
city while Mungishpur is 28 kms away. Mungishpur represents drought prone
area which is favourable rainfed during years of high rainfall while Chandpur
represents shallow and submergence prone area which is favourable rainfed during
the years of low rainfall. In these sites, on-farm experiments on component
technologies e.g. varietal trials with a farming systems perspective are being
conducted by the research staff of NDUAT, in collaboration with the International
Rice Research Institute under the ICAR-IRRI Collaborative Rainfed Rice Program
funded by the Ford Foundation and International Fund for Agricultural
Development (IFAD).
We interviewed the principal heads (male and female) of 94 households
in Chandpur and 78 households in Mungishpur from 1992- 1993 after which
we classified them by social status (higher, scheduled caste and backward class).
208
Fig. 1.
Illiteracy rates of principal male and female farmers in rainfed villages of Faizabad, Uttar
Pradesh, India.
209
needed for agricultural and household tasks as well as for domestic chores such as
cooking and looking after young siblings especially when the mother has to work
in the fields. Parents are willing to forego the labour of sons and even incur cash
expenses on their education in anticipation of huge dowery in future (earnings and
security for the family) since the son will remain with the family even after marriage
and contribute to household earnings (Bennett, 1989).
In terms of access to agricultural information, women obtain information
about crop cultivation from their husbands, neighbours and other women. Despite
the important roles they play in agriculture, scientists include only male farmers
when conducting on-farm trials and only recently have begun to include women
farmers. Women also perceived themselves as housewives rather than farmers.
Cropping systems
In both villages rice followed by wheat is grown mainly for home
consumption. Land preparation for rice is done in June, transplanting or
broadcasting in July, weeding in August, harvesting in October and threshing in
November. Wheat is sown in January, harvested in March and threshed in April. In
June, land is again prepared for rice. Some farmers mix wheat and mustard and
grow lentils, grams and peas after rice. Sugarcane is also grown in the upland areas
from February to May. The availability of tubewells (8 in Chandpur and 3 in
Mungishpur) enables few farmers in Chandpur to grow vegetables which are sold
in the market.
Rice area and yields
Fig. 2 shows the average rice area holding across villages and by
socioeconomic status. Landholding devoted to rice is higher in Mungishpur than in
Fig. 2.
210
Average rice area by social class in rainfed villages of Faizabad, Uttar Pradesh, India.
nearest urban
cattle for milk
diversification
increasing rice
In general, higher castes have larger rice farms (0.97 ha in Chandpur and 1.37
ha in Mungishpur) while the lower castes have smaller areas (0.5 ha and less).
Despite the smaller holding size in Chandpur, the average rice yields are
higher (2.0 to 3.24 tons/ha) compared with Mungishpur (less than 2 tons/ha). The
farmers from the higher castes obtained higher yields per hectare than the farmers
from the lower castes (Fig. 3). Richer farmers not only make more profit because
they are "receptive" to new rice technologies but they have cash and capital to
purchase for "input intensive" system. On the other hand, small farmers are less
willing to take the risks of applying inputs as their lands are often located in less
favourable conditions.
Fig. 3.
Average rice yield by social class in rainfed villages of Faizabad, Uttar Pradesh, India
U.P. is once in four years. Percentage occurrence of droughts of moderate class and
worse in the Kharif season is in the range of 20-25% in eastern U.P.. In addition, long
breaks in rainy days is a common phenomenon in India even during normal years.
Such breaks are common in U.P. and they result in contingent and invisible droughts.
Due to the extreme variability in the climatic conditions, farmers, particularly
in the drought prone areas are not willing to invest more inputs such as fertilizer
and herbicides.
Furthermore, the caste system which creates
distribution of resources is prevalent in eastern U.P.
Fig. 4.
212
Total labour use in rice farming by gender and social class in Chandpur and Mungishpur,
Faizabad, Uttar Pradesh India.
for fuel. In contrast, female family members among the lower castes work on their
farms as well as in other farms as exchange or hired labourers. In Mungishpur,
exchange labour is still widely practiced as a strategy for the lack of cash to pay for
hired labourers. Fig.4 shows higher labour input of female adult workers in rice
cultivation both in higher and lower caste farming families.
Table 1.
Activity
Prepare land
Apply manure/
chemical
Irrigate
Broadcast
Transplant
Weed
Harvest
Thresh
Totaln=94
Both
3.23
3.63
0.00
2.29
3.23
5.92
1.52
3.30
2.05
1.55
5.62
1.04
21.94
0.27
2.41
7.29
13.42
10.83
17.16
53.67
% contribution
Table 2.
Hired
30
70
Adult
Adult
Male Female
Both
0.95
0.65
0.00
0.12
0.95
0.77
1.79
5.71
9.34
14.97
16.45
18.20
75.61
2.44
0.80
1.64
0.12
1.07
0.15
7.82
0.00
0.06
40.93
33.02
15.20
14.81
104.14
100
Both
4.18
4.28
0.00
2.41
4.18
6.69
2.44
0.86
42.57
33.14
16.27
14.96
111.96
3.96
4.10
3.69
1.67
6.69
1.19
29.76
0.27
2.47
48.22
46.44
26.03
31.97
157.81
4.23
6.57
51.91
48.11
3272
33.16
187.57
100
16
84
100
Hired
Exchange
Total
Adult Adult
Adult Adult
Adult Adult
Adult Adult
Male Female Both Male Female Both Male Female Both Male Female Both
Prepare land
Apply manure/
chemical
Irrigate
Broadcast
Transplant
6.66
3.27
0.00
2.52
2.59
2.97
3.60
1.06
1.99
Weed
Harvest
Thresh
Totat n=78
3.12
4.18
2.85
% contribution
93
Adult
Adult
Male Female
Activity
Total
29.24
36
6.66
5.79
1.56
0.60
3.65
4.96
8.66 12.26
12.22 15.34
10.53 14.71
0.10
0.55
4.66
2.17
2.06
14.28 17.13
51.26 80.50
1.31
13.01
64
100
29
0.00
0.00
1.56
0.60
0.00
0.00
0.00
0.00
0.00
0.00
8.22
3.87
0.05 0.15
0.13 0.68
12.07 16.73
7.86 10.03
0.00
0.00
0.50
0.00
0.00
2.16
1.54
0.00
0.00
266
1.74
269
3.52
8.76
5.49
1.81
0.98
2.16 6.59
1.13 4.31
7.69 43.45
7.05 9.11
4.15 5.46
31.31 44.32
71
100
0.20
0.35
0.15
1.20
16
6.49
84
100
33
0.00
2.52
8.22
6.39
1.11 3.80
2.12
22.89
21.62
19.39
5.64
31.65
27.11
25.98
19.41 23.72
89.06 132.51
67
100
213
Fig. 5.
Labour participation by gender in different rice farming activities, Chandpur and Mungishpur,
Faizabad, War Pradesh, India.
215
Fig. 6.
Average annual off-farm and non-farm income of adult male workers by social class in
Faizabad, Uttar Pradesh, India.
Animal systems
In both villages, animals constitute an integral part of a mixed farming
systems. Large animals (bullock and cattle) provide draft power for land preparation,
manure for crops household fuel and milk for home consumption and income.
Ownership of livestock allows farmers to diversify against risks. In Chandpur, more
than 50% of the total households interviewed have cattle and less than 38% have
bullocks. Because of the proximity of this village to the market, farmers particularly
the backward castes (Yadavs), raise cattle for milk which provides them an additional
source of income. In contrast, more households in Mungishpur raise bullocks than
cattle (Fig. 7). Due to the lower percentage of households which own bullocks in
Chandpur, less family male labour, and more income to hire tractors, more farmers
in this village use tractors for land preparation. In Chandpur, 41%, 18% and 9% of
the backward, scheduled and higher caste households, respectively use tractors. In
contrast, in Mungishpur, 15%, 13% and 8% of the backward, scheduled and higher
castes, respectively, use tractors (Fig.8).
216
Fig. 7.
Percent of households with bullocks and cattles in rainfed villages of Faizabad, Uttar Pradesh,
India.
Fig. 8.
Percentage of households using tractors in rainfed villages of Faizabad, Uttar Pradesh, India.
217
At both sites, animals are fed through cut and carry method due to lack of
common grazing lands. Farmers use crop residues, grasses and weeds from rice
fields to feed their animals. In Mungishpur, farmers grow Chari and berseem
in small plots. During the summer and part of the Kharif season (MarchOctober), rice and wheat straw are fed to the animals. One animal requires about
6 kgs of straw and 6 kgs of green fodder per day. Green fodder is most scarce
from April/May to mid June and available from August to September. Thus, a
few farmers grow berseem from October to March while Chari is grown from
March to August/September at about 0.03 ha of land. In Chandpur, very few
farmers have sufficient homesteads or field plots for fodder production, thus a
few of them rent government owned land to grow berseem or buy foddgr from
other farmers. Female family members are mainly responsible in collecting
grasses and weeds from distant places. With limited grazing lands, women have
to spend half a day daily to collect grasses and weeds, particularly in Chandpur
where there are more milch animals which are predominantly cared by female
family members. Twice a day they will feed and milk the animals, also clean the
shed or area where they are tied and collect the dung to make into fuel cakes.
During the drought periods, the only source of fodder and fuelwood is distant
forests, thus, women walk farther away from their homesteads to collect animal
fodder and fuelwood. With increasing deforestation and the degradation of the
common land, the burden falls on women and female children, whose time and
energy in gathering firewood has increased substantially.
Social consequences of stressed environments
What emerges from this initial assessment is the need for agricultural scientists
to understand not only the consequences of stress environments on rice yields, but
also the social consequences of drought on people especially on women who carry
the burden of fulfilling their multiple roles as a farmer, farm manager, wage labourer,
housewife and mother. With low crop productivity due to the fragile environments,
migration of men to the cities in search of work will continue to increase. The number
of female de facto household heads will also increase and will take over the
management of the rice farms. Increasing the demand for female labour in rice
production, processing, animal care, collection of animal fodder and fuelwood may
lead to changes in cooking habits (fewer meals), less time devoted to child care and
breast feeding. During periods of drought, the problem of securing food for the family
and fodder for the animals are hardships shouldered by women.
The goal of increasing rice productivity will be difficult to achieve if
management, particularly under stressed environments which require new
knowledge and skills, will be left to female family members, the majority of whom
are illiterate and without access to training and extension services. We also expect
that their work burden will increase as they face competing demands on their time
between households duties (caring for their children, collecting water, collecting
fuelwood, etc.) and agricultural responsibilities (management of crops, animals,
collecting animal fodder, processing, etc.) unless their male counterparts or children
share in that activities. The greater time spent outside the homes may have
218
detrimental effects on the welfare of the children unless women receive income
which they can use to provide the basic needs of their children.
The preliminary results of this study reveal that the environmental stress
which cause low crop productivity and income pushes men to engage more of their
time and attention in non-farm jobs and consequently, womens work in rice
cultivation increases. The low crop productivity will further exacerbate the disparity
between the higher and lower castes and women will further be burdened. Women
should be given access to education, training and extension services to enhance their
role in decision-making and in increasing farm productivity. They should be given
knowledge and skills on the timeliness of weeding and transplanting operations
which affect rice yields. Since they are the preservers of seeds, their indigenous
knowledge and varietal preferences are important in rice breeding programmes.
CONCLUSION
a)
The sexual divison of labor is related to the social position of the household
and access to non-farm income.
b)
c)
Farmers can bear more risks in rice cultivation if they have access to income
from non-farm sources and would be receptive to input-intensive
technologies.
b)
Include women farmers in evaluating new rice varieties : In both villages, improved
rice varieties such as NDR-423, NDR- 402, NDR-4001, Mahsuri which are
recommended for these rainfed environments have been distributed and
planted and are currently being evaluated by women cooperators.
c)
d)
Introduce new species of animal fodder : Seedlings of new species of forage have
been introduced to increase the local supply of animal feeds and should be
evaluated by male and female farmers.
e)
219
REFERENCES
Bennett Lynn (1989) Gender and poverty in Asia: Issues and Opportunities Concerning Women in the
Indian Economy. The World Bank, Washington, D.C. USA
Boserup Ester (1970) Womans role in economic development. St. Martins Press, New York, U.S.A..
Singh V P (1992) Rainfed lowland rice-based farming system in eastern India. Paper presented at the
Rainfed Lowland Rice Farming Systems Meeting, Yangoon, Myanmar.
220
SUMMARY
Over the last two decades rice production in Eastern India grew at a
satisfactory rate of about 2.7 percent per year. But the growth was uneven across
states and districts particularly in Bihar, Orissa and Assam and Eastern Madhya
Pradesh where rice production failed to keep up with the population and income
induced growth in demand. Production performance was variable even within
homogeneous agroecological zones. Performance has been particularly poor in the
subhumid plateau regions of Chhotanagpur and Chhatisgarh, which also
experienced high rates of rural-urban migration.
Water control and the level of fertilizer use, which are highly correlated,
were important factors behind the spatial variation in productivity growth. The
level and the growth in rice yield was found to be negatively related with the
amount of rainfall, which suggests poor drainage and waterlogging as a
significant constraint to growth in rice production. Land tenure is a special social
problem as districts with higher incidence of tenancy and landlessness had lower
rice yields and slower growth. An inverse relationship is found between the size
of farm and the level and growth in rice yield, suggesting that the small size of
farm in the region is not a serious constraint to the adoption of input-intensive
technology. The size of farm, the incidence of irrigation and the instability in rice
yield are found important factors behind the spatial variation in fertilizer use
and the growth in rice yield.
INTRODUCTION
In the eastern part of India, rice is the principal staple food and agriculture is
dominated by the rice-based cropping systems. This geographical area comprises of
Assam and the small Northeastern States, West Bengal, Bihar, Orisa and the eastern part
of Madhya Pradesh (MT) and Uttar Pradesh (UP) and is inhabited by nearly 500 million
people. The region accounts for two-third of total rice area in India, but because of lower
productivity, contributes only half to the Indian rice production. Unlike other parts of
India where rice is grown under irrigated conditions, rice farmers of the region are
of rice from regional production increased by about 11 percent over the last two
decades. The growth in the productivity of land in rice cultivation (yield rate) was
2.3 percent per year, similar & the performance of Asia as a whole during this period
(Hossain and Laborte, 1994). But the absolute level of yield was only 2.2 tons per
ha, compared to 5.8 tons in China, 4.5 tons in Indonesia and 2.8 tons in Bangladesh
where the production conditions are similar. Within India, the rice yield has reached
4.8 tons in Punjab (in the North) and 4.6 ton in Tamil Nadu (in the south), and in
several districts in these two States farmers get more than 5.5 tons per hectare (GOI,
1993).
The production performance was also uneven across the states in Eastern
India (Table 1). The growth in production was impressive in Uttar Pradesh,
satisfactory in West Bengal, but dismal in Bihar, Orissa and Assam. The annual
growth in rice production was about 5.3 percent in eastern UP, and 2.7 percent in
West Bengal. Nearly four-fifths of the growth was due to the increase in rice yields
made possible through the diffusion of the modern high yielding varieties. In
Madhya Pradesh rice production barely managed to keep pace with the growth of
population; respectively 2.3 and 2.4 percent per year. In Bihar and Orissa, the growth
in rice production was only about 1.6 percent per year. In Orissa, there was an
absolute decline in the area harvested with rice, while rice yield increased by 1.9
percent per year.
Since the land frontier has been closing, the future growth in agricultural
production in the region must come from the growth in productivity of land. The
growth in rice yield was higher than population growth only in Uttar Pradesh and
West Bengal. In other states the productivity growth was less than two percent.
Assam with a growth in yield at less than one percent per year had the most dismal
performance in raising productivity.
An important characteristic of the rice sector in Eastern India is a large year
to year fluctuation in rice production due to unreliable monsoons. For some years
the production shortfall can be astronomical leading to severe food insecurity for
the small and marginal farmers. For example, in Bihar the production shortfall was
67 percent in 1966, 35 percent in 1979, and 24 percent in 1987 compared to the
preceding year. In Madhya Pradesh, the harvest failure was 50 percent in 1965, 58
percent in 1979 and 38 percent in 1986 and 1987. Farmers may suffer from both
floods and droughts in the same season because of heavy early season rains and
prolonged droughts during the end of the monsoon. This makes rice cultivation a
risky venture for majority of the farmers who operate at a subsistence level.
Eastern India experienced considerable rural-urban migration of population
(Table 1). The average population growth rate over the last two decades was 2.2 percent
per year, while the rural population grew at 1.8 percent. The information presented in
Table 1 suggests that the rural-urban migration is partly related to the performance of
the rice sector. The growth in rural population was almost the same as the total
population in UP and West Bengal which experienced an impressive growth in rice
production. In Bihar and Orissa which had sluggish growth, 0.3 to 0.4 percent of the
people migrated every year to urban areas over the last two decades. The rate of
223
States
Area
Yield
Population
growth
Production
Rural
Total
Assam
0.94
0.92
1.86
n.a.
2.1
West Bengal
0.45
2.27
2.72
2.1
2.2
Bihar
0.04
1.55
1.59
1.8
2.1
-0.34
1.88
1.54
1.4
1.8
0.62
1.68
2.30
1.5
2.4
Uttar Pradesh
1.03
4.28
5.31
1.9
2.1
Eastern India
0.44
2.27
2.71
1.84
2.2
Orissa
Madhya
Pradesh
Source :
Note
The growth rates are obtained by estimating semi- logarithmic trend lines on the time
series data.
Identification
of the region
9
Agroecological region
Northern Subhhumid
with alluvial soils
Land area
(million ha)
Plains
12.2
12
13.2
13
27.8
14
16
9.3
11.7
The location and characteristics of the zones can be seen from Figs. 2 to Fig.
6, which are reproduced from Sehgal et al. (1990).
Table 2 reports the estimates of growth rates in cereal production, and the
sources of growth in the production of rice for the five agroecological regions. The
production performance is impressive for the three AEZs in the subhumid and
humid plains (the Gangetic plain), which increased cereal production at or above
3.5 percent per year over the 1970-87 period. The production of non-rice crops
increased at a faster rate than rice, but the growth in rice production itself surpassed
population growth.
225
Fig. 1.
the Eastern humid plains, and Northern subhumid plains had yields over 2.0 tons
per hectare, while most of the districts in the other three AEZs had yields below
that level. The important point to note is that there is a large variation in the level
of yield across districts in all five agroecological zones.
Table 2.
Major
Rice
by
Cereals
production
agroecoregions
area
Yield
Production
0.72
(0.29)
2.68
(0.97)
3.43
(1.08)
4.32
(0.62)
12
0.54
(0.05
1.68
(1.11)
2.22
(1.13)
2.24
(1.06)
13
-0.28
(0.20)
1.21
(0.65)
0.92
(0.81)
1.03
(0.75)
14
0.72
(0.19)
2.05
(0.47)
2.78
(0.53)
3.47
(0.36)
16
0.45
(0.19)
2.76
(0.40)
3.22
(0.51)
3.45
(0.48)
Note:
The performance with respect to the growth of rice yield also varied widely
across districts. In 19 of the sixty districts under study, the growth in rice yield was
more than 3.0 percent per year; while in nine districts the growth rate was negative,
and most of them are located in the Chhotanagpur plateau. Again, the inter-district
variation in the growth of rice yield was large for each of the AEZ's.
227
Fig. 2.
228
Fig. 3.
229
Fig. 4.
230
Fig. 5.
231
Fig. 6.
232
We conclude from the findings that agroecological factors alone cannot explain
the diverse production performance of the rice sector across districts in Eastern India.
Agroecological
zones
< 1.5
1.52.0
2.03.0
3.0 &
over
Total
10
8
10
3
3
3
5
12
10
17
28.3
23
38.3
18
60
30.3
3.3
100.0
20
The socioeconomic factors considered in the analysis are the average size of
the farm in the district, the pattern of distribution of land (measured by the share
of landless agricultural laborers and tenants in total farm population), the incidence
of poverty (measured by the per capita calorie intake of the population for the base
period), and the economic capacity of the farmers (measured by the amount of
chemical fertilizer used per hectare of land in the district). The smaller the size of
farm, the higher the pressure of population on land and hence higher would be the
need to increase foodgrain production. But the size of farm also determines the
availability of labour input and its opportunity cost to the farm household. Small
farms may use more labor and hence achieve higher levels of productivity than large
farms (Berry and Cline, 1979). Small farms may however be economically unable to
afford relatively capital intensive modern technology, and hence the growth in rice
yields may be slower in districts with smaller farm size. The effect of farm size on
productivity would depend on the relative strength of these two oppositive forces.
The unequal distribution of land ownership and the high incidence of tenancy may
provide disincentives to production, and hence is expected to have a negative impact
233
on the growth of rice yield. The subsistence pressure on the growth of rice production
is expected to be captured by the incidence of poverty at the base period.
Table 4.
Agroecological
zones
Up to
1.0
1-0
2.0
2.03.0
Over
3.0
Total
10
20
12
10
11
12
15
13
60
15.0
18.3
20.0
25.0
21.7
100.0
The regression coefficients were estimated using the ordinary Leasts square
method. The results are reported in Table 5.
Table 5.
Factors affecting variation in the level and growth of rice yields across
districts: Regression estimates.
Rice yield
(kg/ha)
Explanatory variables
1
Farm size (ha)
Incidence of landlessness and tenancy
(ratio of total workers)
Per capita energy intake at base
period (10 2 calorie)
Fertilizer use
(kg of materials/ha)
Area under irrigation
(ratio of total area)
Area under upland rice
(ratio of total area)
Eq(1)
Eq(2)
Eq(3)
Eq(4)
-100
(1.15)
-103
(-1.10)
-0.61
(-1.66)
-0.62
(-1.62)
-73
(-0.20)
121
(1.32)
-96
(-0.25)
-3.67**
(-2.37)
-3.11**
(1.95)
77
(0.79)
-0.37
(-0.95)
-0.51
(1.29)
3.3**
(2.97)
855**
(3.26)
314
(0.65)
0.011**
(2.32)
1183**
(4.62)
0.95
(0.86)
2.04*
(1.94)
272
(0.52)
6.78**
(3.30)
6.64**
(3.10)
-3.62**
(-2.76)
46.6**
(-3.41)
0.15**
(-2.72)
-0.19**
(-3.32)
-814**
(-3.92)
-1054**
(-5.12)
-0.41
(-0.57)
-1.290
(-1.42)
-909**
(-5.20)
-1420**
(-6.79)
-1.30
(-1.76)
-2.07**
(-3.08)
Chhotanagpur Plateau
-510**
(-2.46)
-867**
(-4.79)
-0.82
(-0.94)
-2.00**
(2.69)
Chhatisgarh Plateau
-620**
(2.63)
-976**
(-4.45)
-0.54
(-0.54)
-1.72*
(-1.92)
Constant term
2.26**
(5.77)
2.82**
(7.60)
5.29*
(3.19)
7.12
(4.69)
0.77
0.73
0.61
0.56
R2
Note
Figures in parentheses are estimated 't' values. ** denotes that the regression coefficient is
statistically significant at 5% probability level and l at 10 percent.
235
The rice yield is found to be negatively correlated with the amount of rainfall.
The districts with higher rainfall are also found to have slower growth in rice yield.
The regression coefficient for this variable is statistically highly significant. The
results suggest that poor drainage and flooding is a major constraint to increase in
rice yield in Eastern India.
As expected, irrigation is an important factor affecting rice production. The
conversion of rainfed to irrigated land is found to increase rice yield by about 1.2
tons per hectare. Irrigation contributes to the growth in rice yield partly through
facilitating adoption of modern high yielding varieties and larger use of chemical
fertilizers. When the fertilizer variable is included in the model along with the
irrigation variable, the value of the regression coefficient for irrigation is substantially
reduced (Table 5). In fact, in the equation on growth of rice yield, the irrigation
coefficient becomes statistically insignificant as the fertilizer variable picks up the
effect of irrigation. The value of the regression coefficient of fertilizer suggests that
at the farmers level, one kg of additional use of fertilizer (materials) increases rice
production on the margin by 3.3 kg.
Other factors unchanged, the average rice yield is found to be higher in
districts with larger proportion of area under uplands. The positive and statistically
significant regression coefficient for this variable in equations 3 and 4 (in Table 5)
suggests that these districts achieved higher rates of growth in rice yield than the
districts with larger proportion of rainfed lowlands. This finding is contrary to the
prevailing notion that the progress in increasing rice productivity has been extremely
limited in the uplands. This issue needs further investigation.
The association between the level of rice yield and the per capita energy intake
is positive but statistically insignificant. The positive relationship suggests that the
causality may be working the other way - the higher the rice yield, the higher the
per capita energy intake, ie., the lower the incidence of poverty. This result signifies
the importance of increasing rice yield for achieving food secutity. However, the
districts with higher levels of food security have achieved slower growth in rice
yields, as shown by the negative coefficient of the energy intake variable in the
equations for the growth in rice yields.
The districts with smaller size of farms have achieved higher levels of rice
yields and also have higher rates of growth in yields. This is contrary to the notion
that small farms cannot particitate in the adoption of input-intensive technology as
much as the large farms (Griffin, 1974; Pears, 1980). The finding rather supports the
inverse farm size-productivity relationship documented by large number of
micro-studies for South Asia.
The regression coefficient of the landlessness and tenancy variables is negative
and statistically significant in the equation for the growth in rice yields. The value
of the coefficient suggests that the districts with more skewed distribution of land
have lower rice yields and had slower growth in yields than the districts with
relatively less unequal land distribution. This finding points to the importance of
land reforms for providing incentives to growth in agricultural production.
236
The dummy variables for the agroecoregions are statistically highly significant
in the equations for the level of rice yield. The results suggest that there are many
other bio-physical variables which could not be included in the analysis have had
impact on rice yield. Compared to the Humid Plains (mostly in West Bengal) rice
yield are substantially lower not only in the Chhotanagpur and Chhatisgarh Plateau,
but also in the Subhumid Plains in the Upper Ganges Delta. The growth in rice yield
was significantly lower in the two plateau regions and also in Eastern Subhumid
Plains compared to the Humid Plains, even after accounting for differences in
rainfall, irrigation, land levels and the land tenure situation, (eq (4) in Table 5). But
when fertilizer variable is included, the value of the coefficients of the regional
dummy variables declines and becomes statistically insignificant (eq 3). Presumably,
the negative effect of the biophysical constraints in these agroecoregions is partly
offset by larger use of fertilizer input.
What are the factors that influence the use of chemical fertilizers? To address
this issue we conducted another multi- variate analysis relating fertilizer use across
districts to some of the agroecological and socioeconomic variables mentioned above.
We added the standard error of the estimated growth in yield as another explanatory
variable to represent a measure of risk involved in rice cultivation which is a common
problem in Eastern India. The results can be reviewed from Table 6.
Fertilizer use is found to be highly negatively correlated with the instability
in rice production and the relationship is statistically significant. Districts with
more unstable growth in rice yield have lower levels of use of chemicals fertilizers.
As expected, the relationship between irrigation and fertilizer use is positive and
statistically significant. Better water control reduces the risk in rice cultivation
and encourages farmers to apply more fertilizer and adopt fertilizer responsive
varieties.
The coefficient of the farm size variable is negative. It indicates that districts
with smaller farm size have higher intensity of fertilizer use, contrary to a priori
expectation. When the regional dummy variables are introduced the negative
coefficients of farm size and the instability in rice yield turn out to be statistically
insignificant, presumably due to high levels of collinearity between these two sets
of variables. Thus, the lower levels of fertilizer use in the subhumid plains and
plateau regions compared to the humid plains is partly due to larger size of farms
with low subsistence pressure and the higher risks in rice cultivation (Table 2).
In a recent study on research prioritization for eastern India, major technical
constraints to increase rice production are reported as drought, weeds, yellow stem
borer, acid soils, bacterial leaf blight, lodging, blast, submergence, zinc deficiency
and, army worm in order of importance (Widawsky and O Toole, 1990). The study
estimates that these top 10 problem areas account for 60 per cent of the losses due
to technical constraints that contribute to nearly 45 per cent of the gap between the
average farm level yield and the yield obtained from on-farm experiments under
optimal environmental conditions. The study on agroecological zoning for India
mentions flooding, poor drainage and water-logging as key constraints to
agricultural production in the humid and subhumid plains in the Gangetic Delta;
237
and drought, problem soils and soil erosion as key constraints in the Chhotanagpur
and Chhatisgarh plateau (Sehgal et al., 1990).
Table 6.
Factors
Eq(2)
-36.15**
(-3.79)
-8.04
(-0.74)
-36.17**
(-2.47)
-13.15
(-0.90)
-1.00
(0.58)
-3.77*
(2.34)
97.13**
(3.33)
83.95**
(3.08)
33.78
(0.50)
-46.12
(-0.77)
Subhumid
Subhumid
Chhotanagpur
Chhatisgarh
Constant
R2
Note
-72.0**
(2.93)
Plains
-68.9**
(3.26)
Plains
-112.2**
(5.08)
Plateau
-103.8**
(-3.52)
Plateau
term
140.36**
(3.30)
207.9**
(5.22)
0.56
0.71
Figures within parentheses are estimated 't' value of the regression coefficents. ** denotes
statistical significance at 5 percent level.
subhumid plains which suffer frequent late season moisture stress. Submergence
tolerance is crucial not only for areas with problems of flooding and poor drainage
(in Ganges Delta) but also for the Chhatisgarh and Chhotanagpur plateau where
rainfall suddenly picks up with frequent heavy rains for a number of consecutive
days during the months of July and August and then the monsoon abruptly ends
in the month of September (See Figs. 3 and 4). Elongating varieties tend to lodge
after the flood water recede. So, semi-dwarf varieties with abilities to short- term
submergence could prove beneficial to large areas of Eastern India.
Development of resistant cultivars to prolonged moisture stress and short
period submergence can stabilize yields and reduce the risk in rice cultivation, and
thereby induce low-income farmers to adopt input-intensive technologies. The
rainfed Lowland Rice Research Consortium is on the right track in emphasizing
stress physiology as the priority area of research for this region.
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